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Refractive state and accommodation in the eyes of free-swimming versus restrained juvenile lemon sharks (Negaprion brevirostris).

Optical measurements of the refractive state of the eyes of various shark species typically have depicted sharks as hyperopic (far-sighted) with little evidence of accommodation (i.e. the ability to change focus for visualizing objects at different distances from the eye). In this study, we used infrared video retinoscopy to measure the refractive state in juvenile lemon sharks (Negaprion brevirostris). This technique allows dynamic measurement of refractive state in free-swimming animals as they pass by an aquarium window. We found that unrestrained lemon sharks are focused emmetropically relative to a 1-m distant photorefractor for the lateral visual field. However, when restrained either right side up or upside down (the latter inducing tonic immobility), the sharks become increasingly hyperopic, an artifact also reported in some other vertebrates. In addition, unrestrained lemon sharks display small amplitude accommodative excursions. Thus, refractive state measurements on restrained sharks in general may not reflect the natural, resting state of the shark eye, but rather, an induced hyperopia and lack of accommodative function. Such an artifact may be present in other vertebrate species, underscoring the need to obtain measurements of refractive state in unrestrained animals.

Accommodation, Ocular↗

Are corresponding points fixed?

Several investigators have claimed that the retinal coordinates of corresponding points shift with vergence eye movements. Two kinds of shifts have been reported. First, global shifts that increase with retinal eccentricity; such shifts would cause a flattening of the horopter at all viewing distances and would facilitate fusion of flat surfaces. Second, local shifts that are centered on the fovea; such shifts would cause a dimple in the horopter near fixation and would facilitate fusion of points fixated at extreme viewing distances. Nearly all of the empirical evidence supporting shifts of corresponding points comes from horopter measurements and from comparisons of subjective and objective fixation disparity. In both cases, the experimenter must infer the retinal coordinates of corresponding points from external measurements. We describe four factors that could affect this inference: (1) changes in the projection from object to image points that accompany eye rotation and accommodation, (2) fixation errors during the experimental measurements, (3) non-uniform retinal stretching, and (4) changes in the perceived direction of a monocular point when presented adjacent to a binocular point. We conducted two experiments that eliminated or compensated for these potential errors. In the first experiment, observers aligned dichoptic test lines using an apparatus and procedure that eliminated all but the third error. In the second experiment, observers judged the alignment of dichoptic afterimages, and this technique eliminates all the errors. The results from both experiments show that the retinal coordinates of corresponding points do not change with vergence eye movements. We conclude that corresponding points are in fixed retinal positions for observers with normal retinal correspondence.

Accommodation, Ocular↗

Plasticity of convergence-dependent variations of cyclovergence with vertical gaze.

Binocular alignment of foveal images is facilitated by cross-couplings of vergence eye movements with distance and direction of gaze. These couplings reduce horizontal, vertical and cyclodisparities at the fovea without using feedback from retinal image disparity. Horizontal vergence is coupled with accommodation. Vertical vergence that aligns tertiary targets in asymmetric convergence is thought to be coupled with convergence and horizontal gaze. Cyclovergence aligns the horizontal retinal meridians during gaze elevation in symmetrical convergence and is coupled with convergence and vertical gaze. The latter vergence-dependent changes of cyclovergence have been described in terms of the orientation of Listing's plane and have been referred to as the binocular extension of Listing's law. Can these couplings be modified? Plasticity has been demonstrated previously for two of the three dimensions of vergence (horizontal and vertical). The current study demonstrates that convergence-dependent changes of the orientation of Listing's plane can be adapted to either exaggerate or to reduce the cyclovergence that normally facilitates alignment of the horizontal meridians of the retinas with one another during gaze elevation in symmetrical convergence. The adaptability of cyclovergence demonstrates a neural mechanism that, in conjunction with the passive forces determined by biomechanical properties of the orbit, could play an active role in implementing Listing's extended law and provide a means for calibrating binocular eye alignment in three dimensions.

Accommodation, Ocular↗

Dynamic accommodation in rhesus monkeys.

The dynamics of Edinger-Westphal (EW) stimulated accommodation were studied in two young rhesus monkeys to understand the relationships between accommodative amplitude and rates of accommodation and disaccommodation. Accommodative responses were recorded with infrared photorefraction at five different amplitudes spanning the full EW stimulated accommodative range available to each eye. Combined exponential and polynomial functions were fit to the accommodation and disaccommodation responses. Derivatives of these functions provided the maximum rates of accommodation and disaccommodation as well as time constants for each amplitude. Maximum rates of EW stimulated accommodation and disaccommodation were found to increase linearly with amplitudes from 0.58 to 17.41 D in the two monkeys. The results suggests that the rate of EW stimulated accommodation is dictated by the amplitude. We conclude that if dynamic accommodative responses are to be compared in monkeys of different ages it is necessary to compare responses for the same accommodative amplitudes in order to draw conclusions about age related changes.

Accommodation, Ocular↗

Blur detection thresholds in childhood myopia: single and dual target presentation.

There is some suggestion that the ability to detect blur may be altered in adults with myopia. Here, we address the question of whether children with myopia have worse blur detection than other children, and whether blur detection in myopic children is related to the rate of myopia progression. We recruited 20 myopes and 20 non-myopes aged between 8 and 12 years. Refractive errors, visual acuity, and contrast sensitivity were measured and the change in refractive error over the past year calculated from clinic records. Blur detection thresholds for two different types of black and white targets (text and scenes), two illumination conditions and two testing protocols were determined using a computer-based forced-choice testing procedure. The two testing protocols used were: (i) dual image presentation where subjects were asked to choose the clearer of the two images, one image always having zero blur, and (ii) single image presentation in which the subject reported whether the image was clear or blurred. Blur discrimination ability under all tested conditions was similar for both refractive error groups. Blur detection thresholds were 0.27+/-0.15 D (myopes) and 0.24+/-0.07 D (non-myopes) for text images. Thresholds were similar when measured with a one log unit reduction in lighting: 0.27+/-0.31 D compared to 0.23+/-0.14 D. Blur detection thresholds were greater for photographic scenes (myopes 0.41+/-0.36 D, non-myopes 0.44+/-0.36 D) and when only a single text image (myopes 0.51+/-0.21 D, non-myopes 0.59+/-0.01 D) was presented, but this increase was measured in both refractive error groups. There was no correlation between blur thresholds and refractive error magnitude, refractive error progression over the past year, or contrast sensitivity. We found that the blur detection ability showed greater individual variation in myopic children. Further work is required to determine whether blur detection ability is of relevance to myopia development.

Accommodation, Ocular↗

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.

Accommodation, Ocular↗

Age-related changes in refractive index distribution and power of the human lens as measured by magnetic resonance micro-imaging in vitro.

We report a new technique for non-invasively mapping the refractive index distribution through the eye lens using magnetic resonance micro-imaging. The technique is applied to map the refractive index distribution throughout the sagittal plane of 18 human eye lenses ranging in age from 14 to 82 years in vitro. The results are compared with standard models for the human eye lens. They confirm that the refractive index distribution, when plotted as a function of normalised lens radius, is a function of lens age and differs both between the equatorial and axial directions and between the anterior and posterior halves of the optical axis. The refractive index of the lens nucleus exhibits a significant reduction with age amounting to 3.4+/-0.6 x 10(-4) years(-1). The contribution of the gradient index (GRIN) to the lens power decreases by 0.286+/-0.067 D/year, accounting almost entirely for the estimated overall change in lens power with age for these lenses, which were probably in their most accommodated state. The results provide experimental verification of hypothesised changes in the GRIN that have previously been invoked as contributing to presbyopia and support the hypothesis that changes in the GRIN are sufficient to offset effects of increasing curvature of human lenses with age in their unaccommodated state.

Accommodation, Ocular↗

Numerical modelling of the accommodating lens.

Data on geometric and material properties of the human lens derived from various published sources are used to construct axisymmetric, large displacement, finite element models of the accommodating lens of subjects aged 11, 29 and 45 years. The nucleus, cortex, capsule and zonule are modelled as linearly elastic materials. The numerical model of the 45-year lens is found to be significantly less effective in accommodating than the 29-year lens, suggesting that the modelling procedure is capable of capturing at least some of the features of presbyopia. The model of the 11-year lens shows some anomalous behaviour, and reasons for this are explored.

Accommodation, Ocular↗

Effects of longitudinal chromatic aberration on accommodation and emmetropization.

PURPOSE: Less accommodation was found when human subjects read in blue (peak at about 440 nm) than when they read in red light (above 600 nm; [Kroger & Binder, British Journal of Ophthalmology 84 (2000) 890]). On the other hand, emmetropization in chickens did not appear to compensate for the chromatic defocus (385 nm versus 665 nm; [Rohrer, Schaeffel & Zrenner, Journal of Physiology 449 (1992) 363]). The apparently contradictory result was studied in more detail in humans and chickens. METHODS: Accommodation was measured with an eccentric infrared photorefractor, the PowerRefractor, in human subjects reading under quasi-monochromatic illumination conditions. Chickens were refracted in quasi-monochromatic ambient illumination but with no particular fixation target. In a second experiment, they were also raised in monochromatic light for two days and subsequently refracted both in complete darkness, in monochromatic light, and in white light, both without and with cycloplegia. RESULTS: Consistent with the initial report by Kroger and Binder [British Journal of Ophthalmology 84 (2000) 890], accommodation in human subjects was found to shift in accordance with the chromatic aberration function. An immediate shift in accommodation tonus was also found in the chickens when they were refracted under red and in blue ambient illumination (average difference between refractions in both conditions: 1.26+/-0.54 D, p<0.001 paired t-test). This value is close to the chromatic focus difference between the two wavelengths (1.5 D [Mandelman & Sivak, Vision Research 23 (1983) 1555]). When chickens were raised in blue or red light for two days, and their refractions were subsequently measured in complete darkness, they showed also a difference in refractions (1.41+/-1.00 D; ANOVA: p<0.0012, post hoc t-test: at least p<0.05 among different groups). This difference was no longer significant when they were refracted in white light but became again significant when they were cyclopleged (0.57+/-0.58 D, p=0.039, unpaired t-test). The latter observation makes it unlikely that the difference resulted just from a shift in the resting tonus of accommodation. CONCLUSIONS: (1) Imposed chromatic defocus produces a shift in accommodation tonus in both humans and chickens which is, in the case of the chicken, followed by a shift in cycloplegic refractive state into the same direction, (2) the difference to the previous study by Rohrer, Schaeffel and Zrenner [Journal of Physiology 449 (1992) 363] can be explained from the fact that shorter wavelengths were used than in the present study, at which emmetropization was no longer functional and, (3) the small amplitude and the variability of the shifts in refraction do not allow clear statements about the role of the "lag of accommodation" in refractive development but they show that several cone types contribute to emmetropization.

Accommodation, Ocular↗

Development of the response AC/A ratio over the first year of life.

This study investigated the development of the link from accommodation to vergence in infants by occluding one eye thus removing binocular cues. Occluded adults continue to converge partially demonstrating that the accommodative drive to vergence (the AC/A link) and proximal cues are sufficient to drive vergence. For infants of all ages, AC/A ratios were found to be in the normal adult range. We conclude that infants can use monocular cues to drive vergence and that this occurs before the age when there is a substantial increase in the accuracy of oculomotor processes. There is flexibility in the developing visual system which is able to produce early vergence responses by relying upon alternative cues.

Accommodation, Ocular↗

An evaluation of the lag of accommodation using photorefraction.

OBJECTIVE: The lag of accommodation which occurs in most human subjects during reading has been proposed to explain the association between reading and myopia. However, the measured lags are variable among different published studies and current knowledge on its magnitude rests largely on measurements with the Canon R-1 autorefractor. Therefore, we have measured it with another technique, eccentric infrared photorefraction (the PowerRefractor), and studied how it can be modified. METHODS: Particular care was taken to ensure correct calibration of the instrument. Ten young adult subjects were refracted both in the fixation axis of the right eye and from the midline between both eyes, while they read text both monocularly and binocularly at 1.5, 2, 3, 4 and 5 D distance ("group 1"). A second group of 10 subjects ("group 2"), measured from the midline between both eyes, was studied to analyze the effects of binocular vs monocular vision, addition of +1 or +2 D lenses, and of letter size. Spherical equivalents (SE) were analyzed in all cases. RESULTS: The lag of accommodation was variable among subjects (standard deviations among groups and viewing distances ranging from 0.18 to 1.07 D) but was significant when the measurements were done in the fixation axis (0.35 D at 3 D target distance to 0.60 D at 5 D with binocular vision; p<0.01 or better all cases). Refracting from the midline between both eyes tended to underestimate the lag of accommodation although this was significant only at 5 D (ANOVA: p<0.0001, post hoc t-test: p<0.05). There was a small improvement in accommodation precision with binocular compared to monocular viewing but significance was reached only for the 5 D reading target (group 1--lags for a 3/4/5 D target: 0.35 vs 0.41 D/0.48 vs 0.47 D/0.60 vs 0.66 D, ANOVA: p<0.0001, post hoc t-test: p<0.05; group 2--0.29 vs 0.12 D, 0.33 vs 0.16 D, 0.23 vs -0.31 D, ANOVA: p<0.0001, post hoc t-test: p<0.05). Adjusting the letter height for constant angular subtense (0.2 deg) induced scarcely more accommodation than keeping letter size constantly at 3.5 mm (ANOVA: p<0.0001, post hoc t-test: n.s.). Positive trial lenses reduced the lag of accommodation under monocular viewing conditions and even reversed it with binocular vision. CONCLUSIONS: After consideration of possible sources of measurement error, the lag of accommodation measured with photorefraction at 3 D (0.41 D SE monocular and 0.35 D SE binocular) was in the range of published values from the Canon R-1 autorefractor. With the measured lag, simulations of the retinal images for a diffraction limited eye suggest surprisingly poor letter contrast on the retina.

Accommodation, Ocular↗

Changes in the internal structure of the human crystalline lens with age and accommodation.

Scheimpflug images were made of the unaccommodated and accommodated right eye of 102 subjects ranging in age between 16 and 65 years. In contrast with earlier Scheimpflug studies, the images were corrected for distortion due to the geometry of the Scheimpflug camera and the refraction of the cornea and the lens itself. The different nuclear and cortical layers of the human crystalline lens were determined using densitometry and it was investigated how the thickness of these layers change with age and accommodation. The results show that, with age, the increase in thickness of the cortex is approximately 7 times greater than that of the nucleus. The increase in thickness of the anterior cortex was found to be 1.5 times greater than that of the posterior cortex. It was also found that specific parts of the cortex, known as C1 and C3, showed no significant change in thickness with age, and that the thickening of the cortex is entirely due to the increase in thickness of the C2 zone. With age, the distance between the sulcus (centre of the nucleus) and the cornea does not change. With accommodation, the nucleus becomes thicker, but the thickness of the cortex remains constant.

Accommodation, Ocular↗

Accommodation in binocular contour rivalry.

Although contour rivalry is known to suppress the contribution of the non-dominant eye to some visuomotor mechanisms such as the pupillary light reflex, there have been no reports of the impact of rivalry on accommodation control. In the situation where the accommodation demands in the two eyes are in dynamic conflict, it has been reported that the accommodation response can be modelled in terms of a vector average of the appropriate response in the two eyes. This study compared the binocular interactions in the accommodation system with rivalrous and non-rivalrous stimuli. Accommodation was continuously monitored with an infrared optometer, while the accommodation demand in the two eyes was dynamically modulated independently in the two eyes. When the visual target was perceptually rivalrous the previously described binocular interactions were abolished and the accommodation response closely followed the accommodation demand presented to the dominant eye.

Accommodation, Ocular↗

Local changes in eye growth induced by imposed local refractive error despite active accommodation.

We have tested whether defocus imposed on local retinal areas can produce local changes in eye growth, even if accommodation is available to clear part of the imposed defocus. Hemi-field lenses were attached to little leather hoods that were worn by young chickens from day 11-15 post-hatching. The lens segments defocused either the nasal or the temporal visual field, or covered the full field. We found that negative lenses (-7.5 D) were incompletely compensated in all three cases but caused significant myopia in the defocused parts of the visual field (differences to fellow eyes with normal vision: nasal visual field -3.13 +/- 1.56 D, P < 0.001; temporal visual field -4.02 +/- 1.38 D, P < 0.001; full field -3.82 +/- 2.48 D, P = 0.01). Myopia was not enhanced if the lenses covered the entire visual field. Positive lenses (+6.9 D) caused larger changes in refraction than negative lenses and, again, there was no significant difference in the amount of induced hyperopia in the nasal or temporal retina, or in the amount of hyperopia with full-field lenses (difference to fellow eyes with normal vision: nasal visual field +6.2 +/- 2.69 D, P < 0.001; temporal visual field +5.95 +/- 2.22 D, full field +7.22 +/- 2.44 D, P < 0.001). To compare the shapes of the excised eyes after lens treatment, we wrote a fully automated image processing program that traced their outlines in digitized video images. We found that the shapes of the eyes treated with positive lenses did scarcely differ from their fellow eyes with normal vision, indicating that hyperopia over this 4 day period was caused mostly by choroidal thickening. Full field negative lenses produced significant axial eye elongation; the effects of locally imposed defocus on eye shape were less conspicuous and were significant only in some areas. That local compensation of defocus was possible for both negative and positive lenses, suggests that the retina can recognize the sign of defocus without accommodation cues. Even more striking is that the presence of accommodation is apparently ignored since the drift in the plane of focus during accommodation does not disturb the compensation process. We re-analyze previous experimental results that argue for different mechanisms for deprivation myopia and lens-induced refractive errors. We propose that lens-induced refractive errors are compensated by similar retinal mechanisms as the ones proposed by Bartmann and Schaeffel [(1994). Vision Research, 34, pp. 873-876] to explain deprivation myopia. The proposed mechanisms can integrate with long time constants over the spatial frequency content in the retinal image while the viewing distances change, and control both choroidal thickening and scleral growth. However, it turns out that the compensation of imposed myopia cannot be explained if only one constant viewing is available. Apparently, there is more than a retinal blur detector to guide refractive development.

Accommodation, Ocular↗

Contrast sensitivity in infants and children with Down syndrome.

A new contrast sensitivity (CS) card test was used to estimate contrast sensitivity in 18 infants and children with Down syndrome (DS). The results showed that although the overall shape of the contrast sensitivity functions (CSFs) of the subjects with DS was the typical inverted-U, their CSFs were depressed in comparison to control subjects and this relative loss became larger with increasing spatial frequency. In addition, there was little improvement in CS with age and the mean CSF among children with DS (mean age = 7.3 years) was equivalent statistically to a group of 12-month-olds without DS. The Teller Acuity Cards (TAC) were also used to assess visual acuity in 17 of the 18 children in our sample. The results of these tests showed that their visual acuity (VA) was significantly lower than normal, but was consistent with that extrapolated from each subject's CSF. Taken together with previous anatomical and developmental findings, our results suggest that the deficits in spatial vision among children with DS is due primarily to restricted cortical development, and secondarily, to the additional accommodative and ocular conditions that are prevalent in this population.

Adolescent↗

Presbyopia and the optical changes in the human crystalline lens with age.

Lenses from 27 human eyes ranging in age from 10 to 87 years were used to determine how accommodation and age affect the optical properties of the lens. A scanning laser technique was used to measure focal length and spherical aberration of the lenses, while the lenses were subjected to stretching forces applied through the ciliary body/zonular complex. The focal length of all unstretched lenses increased linearly with increasing age. Younger lenses were able to undergo significant changes in focal length with stretching, whereas lenses older than 60 years of age showed no changes in focal length with stretching. These data provide additional evidence for predominantly lens-based theories of presbyopia. Further, these results show that there are substantial optical changes in the human lens with increasing age and during accommodation, since both the magnitude and the sign of the spherical aberration change with age and stretching. These results show that the optical properties of the older presbyopic lens are quite different from the younger, accommodated lens.

Accommodation, Ocular↗

Static aspects of accommodation: age and presbyopia.

Although the progressive reduction in accommodative amplitude with increased age is well documented, little is known about several other aspects of static or steady-state accommodation to provide a comprehensive assessment of changes related to age and presbyopia. Static components of accommodation (tonic accommodation, depth-of-focus, slope of the stimulus/response function, and accommodative controller gain) were assessed objectively using an infrared (IR) optometer in 30 human subjects aged 21-50 years; depth-of-focus was also determined psychophysically as was accommodative amplitude. Tonic accommodation and the amplitude of accommodation decreased with increased age, whereas the subjective depth-of-focus increased; the other parameters remained unchanged. The decrease in tonic accommodation and amplitude of accommodation was attributed to biomechanical factors, whereas the increase in subjective depth-of-focus was believed to result from increased tolerance to defocus related to the gradual onset of presbyopia. Constancy of the objective depth-of-focus suggested absence of age effects on the neurologic control of reflex accommodation, whereas the lack of systematic change in slope and controller gain provided support for the Hess-Gullstrand theory of accommodation and presbyopia.

Accommodation, Ocular↗