The absence of significant age-related changes in red-green isoluminance values indicates that optical and receptoral factors do not significantly affect the relative sensitivity to one of these colors selectively. The significant correlation between age and red-blue isoluminance suggests that older observers with phakic eyes require a lower level of red luminance for red to be perceived as having the same luminance as blue. In contrast, observers with pseudophakic eyes obtained results similar to those of the youngest observers (observers younger than 30 years and in their 30s) in the red-blue isoluminance task. This indicates that the age-related decreases in relative sensitivity to short-wavelength light, as assessed by the minimum motion technique, can be accounted for by optical factors. Overall, these results are in agreement with earlier research suggesting that sensitivity to shorter wavelength stimuli is more affected than sensitivity to longer wavelength stimuli.
7 8 9
If age-related changes at the level of the receptors preferentially affecting one type of receptor were the cause of the relative loss of sensitivity to short wavelength light, one would not expect those with pseudophakic eyes to obtain results similar to those of young observers in the red-blue isoluminance task. That no significant age-related changes were found in red-green and red-blue isoluminance values in pseudophakic eyes suggests that there are no selective losses in the relative sensitivity of one cone mechanism. This is in agreement with earlier findings, which showed that all three types of cone lose overall sensitivity at a similar rate.
17
The motion-nulling task found evidence of a small but significant age-related decrease in the sensitivity of the L-M postreceptoral chromatic mechanism in comparison with the luminance mechanism. However, one should be cautious when considering this result, because age accounts for only approximately 10% of the variance among all observers on this task. This age-related decrease in relative sensitivity to chromatic stimuli is not consistent with earlier literature suggesting that age-related losses in the luminance and chromatic pathways are nonselective,
27 although this discrepancy may occur because observer age accounts for only a small proportion of the variance. Because the postreceptoral losses in sensitivity to red-green chromatic motion were relatively small, it is unlikely that they account for much of the selective losses in sensitivity to short wavelength stimuli. Furthermore, it is unlikely that age-related losses in the sensitivity of the L-M postreceptoral system can account for the systematic age-related decrease in the results of our red-blue isoluminance task, because these losses were quite small.
It may be argued that, in the motion-nulling task, the drifting red-green grating did not isolate the L-M red-green postreceptoral mechanism and that there is a contribution of the S-(L+M) blue-yellow mechanism to the perception of chromatic motion.
28 Should this be so, it is also possible that the reported small age-related changes in the red-green motion-nulling task can reflect losses in the S-(L+M) postreceptoral mechanism. The isoluminant red-green grating used in these experiments stimulated both chromatic mechanisms, although it stimulated the L-M system preferentially. Research on chromatic motion studying the S-(L+M) system has found that chromatic gratings isolating this mechanism resulted in lower perceived speeds,
29 30 even when matched for multiples of threshold.
29 In addition, when a chromatic grating stimulates both postreceptoral color mechanisms, perceived speed appears to be determined using a single mechanism.
30 Thus, because the chromatic grating stimulated the L-M mechanism preferentially, the L-M system is more sensitive to contrast, and the S-(L+M) appears to be less efficient at chromatic motion perception,
24 29 30 it is probable that results reflect losses in the L-M chromatic system.
That older observers with pseudophakic eyes obtained results identical with those of the youngest observers on a red-blue isoluminance task suggests that lenticular senescence can account for most, if not all age-related changes found in phakic eyes on this task. To determine whether lenticular senescence alone could explain the age-related decrease in red-blue isoluminance, a model based on lens absorbance as a function of age and wavelength
14 was created.
This model describes the mean luminance of the red and blue phosphors as vectors in a color space. A representation of such a space is described in
Figure 3 .
The length of the
L R and
L B vectors represents the mean luminance of the red and blue phosphors, respectively, on a red-blue isoluminance task. In such a space, it is assumed that isoluminance is achieved when a certain distance is reached between the two vectors (
k). This
k vector is perpendicular to the vector representing blue luminance. This color space is illustrated in
Figure 4 . The value of
k can be determined with the function
\[k\ {=}\ \sqrt{L_{\mathrm{R}}^{2}\ {-}\ D_{\mathrm{B}}^{2}}.\]
If lenticular senescence can account for all the age-related changes on a red-blue isoluminance task for phakic observers, the value of k remains constant with age once this factor is taken into account.
To determine the value of
k, it was necessary to determine the amount of light from the monitor’s blue phosphor reaching the retina. To do this, the spectrum of the blue phosphor of the monitor was filtered using the lens absorbance values of Weale.
14 Because the pupil size of observers was not measured during testing, it is assumed that all observers had a pupil diameter of 4 mm during this period. It should be noted that although pupil size may attenuate light reaching the retina, these effects are not wavelength selective. Therefore, senile miosis by itself should not cause a change in the relative luminances of the red and blue phosphors reaching the retina. Moreover, pupil size also diminishes for observers with pseudophakic eyes, and yet, they obtain results similar to those of the young with phakic eyes on the red-blue isoluminance task. In addition, if pupil size were an important factor on red-blue isoluminance, one would expect age-related changes in these values for observers with pseudophakic eyes. We observed no evidence of this. Because absorbances
14 were available only at wavelengths of 380, 400, 420, 440, and 460 nm, the best-fitting exponential function was used to estimate lens absorbance at intermediate wavelengths for each age. These values were used to determine the transmissivity (
T) of the ocular lens as a function of wavelength and age (
A) with the formula
\[T({\lambda},A)\ {=}\ 10^{{-}D({\lambda},A)}\]
where
D is the absorbance of the ocular lens at a given wavelength and age. Weale’s values of lenticular absorbance
14 indicate that the amount of long-wavelength light absorbed by the crystalline lens at any age is not significant. Because of this, the red phosphor luminance values were not filtered. Reflectometry results show this clearly.
26
These values of lens transmissivity as a function of wavelength were used to estimate the proportion of light from the blue phosphor transmitted by the crystalline lens at different ages using the formula
\[T_{\mathrm{B}}(A)\ {=}\ {{\sum}}T({\lambda},A)\ {\cdot}\ P({\lambda})\]
where
P(λ) is the proportion of the blue phosphor’s output at a given wavelength over the total output of the phosphor. This allowed us to evaluate the transmittance of crystalline lens to the blue phosphor in observers aged 20, 40, 60, and 80 years. These values were then used to estimate the proportion of light from the blue phosphor transmitted by the lens in observers of ages of 30, 50, and 70 years.
The estimated transmissivity of the crystalline lens to the blue phosphor was used to determine the amount of light from the blue phosphor transmitted to the retina at a given age with the function
\[L{^\prime}_{\mathrm{B}}(A)\ {=}\ L_{\mathrm{B}}\ {\cdot}\ T_{\mathrm{B}}(A)\]
where
L′
B(
A) is the filtered mean luminance of the blue phosphor at a given age, once lenticular senescence is taken into account. This filtered value of the blue phosphor’s mean luminance was used to determine
k at the age of 20 with the following equation
\[k\ {=}\ \sqrt{L_{\mathrm{R}}^{2}(20)\ {-}\ L{^\prime}_{\mathrm{B}}^{2}(20)}.\]
To estimate the mean luminance of the red phosphor, which appears isoluminant with the filtered values of blue luminance at this age, the color luminance contrast results on a red-blue isoluminance task was estimated by using the best-fitting regression line in
Figure 2B .
The
k value determined in
equation 10 and the filtered mean luminance of the blue phosphor were used to estimate the predicted mean red luminance (
L′
R) necessary to achieve isoluminance with blue at different ages. This was determined by
\[L{^\prime}_{\mathrm{R}}(A)\ {=}\ \sqrt{L{^\prime}_{\mathrm{B}}^{2}(A){+}k^{2}}.\]
The predicted mean luminance of the red phosphor was used to determine the predicted color luminance contrasts as a function of age with
equation 1 . The predicted color luminance contrasts made by this model and the experimental results are shown in
Figure 5 . As can be seen, the model shows good fit with the best-fitting regression line for red-blue isoluminance results. This suggests that the optical factor of lenticular senescence can account for the age-related decrease in red-blue isoluminance values alone. This is concordant with the finding that observers with pseudophakia performed similarly with younger observers with phakic eyes on a red-blue isoluminance task (see
Fig. 2B ). This is also consistent with earlier suggestions that no selective losses occurred in the various cone mechanisms.
17
That older observers with pseudophakic eyes performed as well as younger observers on the red-blue isoluminance task led us to conclude that lenticular senescence can account for the decrease in sensitivity to short-wavelength visual stimuli observed in the experiments discussed in this article. No evidence was found for a selective loss of one cone type in normal aging. This conclusion is also supported by the fact that a model based on lenticular senescence showed such good fit with the data in the red-blue isoluminance task. Furthermore, our data showed a small but significant decrease in the sensitivity of postreceptoral chromatic mechanism relative to luminance contrast sensitivity. It is not currently possible to determine whether there is a selective loss in one of the two postreceptoral chromatic mechanisms. A motion-nulling task seems ill-suited as a method of comparing losses in these two systems, because the S-(L+M) mechanism does not seem to contribute as much to motion processing.
22 24 25 29 30
The authors thank Vasile Diaconu for helpful discussions.