Homeostasis of Eye Growth and the Question of Myopia

Evolution of eyes

The results of many studies in a variety of species have significantly advanced our understanding of the role of visual experience and the mechanisms of postnatal eye growth, and the development of myopia. This paper surveys and reviews the major contributions that experimental studies using animal models have made to our thinking about emmetropization and development of myopia. These studies established important concepts informing our knowledge of the visual regulation of eye growth and refractive development and have transformed treatment strategies for myopia. Several major findings have come from studies of experimental animal models. These include the eye's ability to detect the sign of retinal defocus and undergo compensatory growth, the local retinal control of eye growth, regulatory changes in choroidal thickness, and the identification of components in the biochemistry of eye growth leading to the characterization of signal cascades regulating eye growth and refractive state. Several of these findings provided the proofs of concepts that form the scientific basis of new and effective clinical treatments for controlling myopia progression in humans. Experimental animal models continue to provide new insights into the cellular and molecular mechanisms of eye growth control, including the identification of potential new targets for drug development and future treatments needed to stem the increasing prevalence of myopia and the vision-threatening conditions associated with this disease.

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IMI - Report on Experimental Models of Emmetropization and Myopia.

A paraxial schematic eye model for the growing C57BL/6 mouse.

Recent studies have shown that certain nocturnal insect and vertebrate species have true color vision under nocturnal illumination. Thus, their vision is potentially affected by changes in the spectral quality of twilight and nocturnal illumination, due to the presence or absence of the moon, artificial light pollution and other factors. We investigated this in the following manner. First we measured the spectral irradiance (from 300 to 700 nm) during the day, sunset, twilight, full moon, new moon, and in the presence of high levels of light pollution. The spectra were then converted to both human-based chromaticities and to relative quantum catches for the nocturnal hawkmoth Deilephila elpenor, which has color vision. The reflectance spectra of various flowers and leaves and the red hindwings of D. elpenor were also converted to chromaticities and relative quantum catches. Finally, the achromatic and chromatic contrasts (with and without von Kries color constancy) of the flowers and hindwings against a leaf background were determined under the various lighting environments. The twilight and nocturnal illuminants were substantially different from each other, resulting in significantly different contrasts. The addition of von Kries color constancy significantly reduced the effect of changing illuminants on chromatic contrast, suggesting that, even in this light-limited environment, the ability of color vision to provide reliable signals under changing illuminants may offset the concurrent threefold decrease in sensitivity and spatial resolution. Given this, color vision may be more common in crepuscular and nocturnal species than previously considered.

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Crepuscular and Nocturnal Illumination and Its Effects on Color Perception by the Nocturnal Hawkmoth Deilephila elpenor

The focal length of the vertebrate eye is a function of wavelength, i.e. the eye suffers from longitudinal chromatic aberration. Chromatic defocus is a particularly severe problem in eyes with high light-gathering ability, since depth of field is small due to a pupillary opening that is large in relation to the focal length of the eye. Calculations show that in such eyes only a narrow spectral band of light can be in focus on the retina. For the major part of the visual spectrum, spatial resolution should be limited by the optics of the eye and far lower than the resolving power achievable by the retinal cone photoreceptor mosaic. To solve this problem, fishes with irises unresponsive to light have developed lenses with multiple focal lengths. Well-focused images are created at the wavelengths of maximum absorbance of all spectral cone types. Multifocal lenses also appear to be present in some terrestrial species. In eyes with mobile irises, multifocal lenses are correlated with pupil shapes that allow all zones of the lens, with different refractive powers, to participate in the imaging process, irrespective of the state of pupil constriction.

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Multifocal lenses compensate for chromatic defocus in vertebrate eyes.

By rearing fish in various monochromatic illuminations we investigated (1) the potential for compensation of refractive error due to chromatic aberration, (2) the contributions of the chromatic channels to emmetropization, and (3) the role of color cues in the control of eye growth. Cichlid fish (Aequidens pulcher) were reared for 6 months (12 h light/12 h dark) in monochromatic lights (623.5, 534.1, 485.0 nm; spectral purity 5–10 nm). Light levels were isoirradiant at 1.1·1012 quanta/s/cm2. Two control groups were reared in white light with down-welling illuminances of 0.2 and 33 lx. Nasotemporal diameters (NTDs) of the eyes were measured in relation to lens size. Due to the oblique axis of highest acuity vision in cichlids, NTD is considered to be a more important dimension than axial length. Variances in NTD were equally small in all rearing groups. NTDs were enlarged with increasing wavelengths of the rearing lights with highly significant values over controls in the red-light group. The wavelength-dependent size of the eyes matched the changes in focal length due to longitudinal chromatic aberration. Complete recovery from eye enlargement was observed after fish reared in red light were exposed to a white light regime for 5 weeks. Small variances in NTD in all groups indicated stringent control of eye growth in the absence of color cues. The reversibility of the increase in NTD in fish reared in red light suggests that the eyes were emmetropized by visually guided mechanisms. Eye size in fish reared in white light was intermediate between the values expected if only blue-sensitive single or the red- and green-sensitive double cones contributed to the control of eye growth. This suggests that all chromatic channels participate in emmetropizing the fish eye.

The eye of the blue acara (Aequidens pulcher, Cichlidae) grows to compensate for defocus due to chromatic aberration.

Vietnamese leaf turtles (Geoemyda spengleri) were tested for their ability to focus on prey objects at various distances. Accommodation was continuously measured by infrared photoretinoscopy. All animals investigated during this study showed a surprisingly high precision of accommodation over a range of over 30 D. Measured accommodation matched the target distance accurately for distances between 3 and 17 cm. The tur- tles switched between independent and coupled accom- modation in the two eyes. Independent accommodation was observed when the turtles inspected their environ- ment visually without a defined object of interest. Cou- pled accommodation was only observed during binocular prey fixation. When a turtle aimed at a target, the symmetrical focus of both eyes persisted even if vision was totally blocked in one eye or altered by ophthalmic lenses. This suggests that the eyes were linked by internal neuronal mechanisms. The pupil of the eye responded clearly to changes in ambient light intensity. A strong decrease in pupil size was also ob- served when the eye was focused on a close target. In this case, the constriction of the pupil probably aids in the deformation of the eye lens during near-accommodation.

H.-J. Wagner

Papers

Multifocal lenses compensate for chromatic defocus in vertebrate eyes.

The eye of the blue acara (Aequidens pulcher, Cichlidae) grows to compensate for defocus due to chromatic aberration.

Accommodation behaviour during prey capture in the vietnamese leaf turtle (Geoemyda spengleri)