Age-related changes in wavelength discrimination.

Wavelength discrimination functions (420 to 620-650 nm) were measured for four younger (mean 30.9 years) and four older (mean 72.5 years) observers. Stimuli consisted of individually determined isoluminant monochromatic lights (10 Td) presented in each half of a 2 degrees circular bipartite field with use of a Maxwellian-view optical system. A spatial two-alternative forced-choice method was used in combination with a staircase procedure to determine discrimination thresholds across the spectrum. Small but consistent elevations in discrimination thresholds were found for older compared with younger observers. Because the retinal illuminance of the stimuli was equated across all observers, these age-related losses in discrimination are attributable to neural changes. Analyses of these data reveal a significant change in Weber fraction across adulthood for a chromatically opponent pathway receiving primarily antagonistic signals from middle-wavelength-sensitive and long-wavelength-sensitive cones but not for a short-wavelength-sensitive cone pathway.

Senescence of foveal and parafoveal cone sensitivities and their relations to macular pigment density.

Foveal and parafoveal increment thresholds were measured for 50 observers (12-88 years of age) under conditions that isolated retinal mechanisms dominated by short- (S-), middle- (M-), or long- (L-) wave-sensitive cones. Thresholds were obtained on the plateau of the threshold-versus-intensity function of each isolated mechanism and were referred to the retina by using individual measurements of ocular media and macular pigment density. Age-related increases in foveal thresholds, specified at the retina, were found for all three cone mechanisms. Parallel sensitivity losses for each cone mechanism were also observed at 4 degrees and 8 degrees in the temporal retina. A significant positive correlation was found between foveal macular pigment density and the S-cone, but not the M- and L-cone, log sensitivity difference (0 degrees-8 degrees) specified at the retina. This relation is expected from the hypothesis that the macular pigment protects the photoreceptors from senescent losses in sensitivity. However, because this result is independent of age, it is interpreted as being due to local gain changes resulting from differential filtering of incident light by the macular pigment between the fovea and the parafovea.

Spatial summation in human cone mechanisms from 0 degrees to 20 degrees in the superior retina.

The maximum area of complete spatial summation (i.e., Ricco's area) for human short-wavelength-sensitive-(S-) and long-wavelength-sensitive- (L-) cone mechanisms was measured psychophysically at the fovea and at 1.5 degrees , 4 degrees , 8 degrees , and 20 degrees along the vertical meridian in the superior retina. Increment thresholds were measured for three observers by a temporal two-alternative forced-choice procedure. Test stimuli ranging from -0.36 to 4.61 log area (min2) were presented on concentric 12.3 degrees adapting and auxiliary fields, which isolated either an S- or an L-cone mechanism on the plateau of its respective threshold versus intensity function. Test flash durations were 50 and 10 ms for the S- and L-cone mechanisms, respectively. The data indicate that, from 0 degrees to 20 degrees, Ricco's area increases monotonically for the L-cone mechanism, is variable for the S-cone mechanism, and is larger for the S-cone mechanism than for the L-cone mechanism for essentially all retinal locations. This pattern of results most likely reflects differences in ganglion cell density and changes in neural convergence with retinal eccentricity.

Senescent changes in scotopic contrast sensitivity.

Scotopic contrast sensitivity functions (CSFs) were measured for 50 observers between the ages of 20 and 88 years. Using a maximum-likelihood, 2-alternative, temporal forced-choice threshold-estimation algorithm, scotopic CSFs were measured at 7 spatial frequencies ranging from 0.2 to 3.0 cpd, with mean retinal illuminance equated for observers at -0.85 log scotopic Trolands. For each stimulus condition, eight cycles of a horizontal sinusoidal grating were presented within +/- 1 S.D. of a 2-D Gaussian-spatial envelope and within a 1-s Gaussian-temporal envelope. Stimuli were centered on the nasal retina along the horizontal meridian 6 degrees from the fovea. Scotopic CSFs were found to be low-pass. Statistically significant age-related declines in contrast sensitivities were found for spatial frequencies at or below 1.2 cpd. There was also a statistically significant decrease in the high frequency cut-off with age (P < 0.01). An explanation of these results in terms of optical factors is rejected, while the results are consistent with age-related changes in the magnocellular pathway.

The area of complete scotopic spatial summation enlarges with age.

The maximal area of complete scotopic spatial summation (Ricco's area) was determined for 50 subjects ranging in age from 19 to 87 yr. Increment thresholds were measured for 10-ms, 520-nm circular test lights of varying diameters that were superimposed and concentric with a 10 degrees, 640-nm circular background. The test lights were imaged in Maxwellian view along the horizontal meridian, 6 degrees nasal from a foveal fixation point. The results demonstrate a statistically significant enlargement of Ricco's area with age. The average angular subtenses of Ricco's areas for the ten youngest (mean = 26 yr) and ten oldest (mean = 75 yr) observers were approximately 48 and 69 arc min, respectively. Model simulations based on a series of optical transfer functions of the eye and varying degrees of intraocular light scatter for younger and older observers show that preneural factors cannot account for these results. Therefore changes in neural mechanisms must be invoked to explain the enlargement in the size of Ricco's area under scotopic conditions.

Spectral mechanisms of spatially induced blackness: data and quantitative model.

Spectral efficiency functions and tests of additivity were obtained with three observers to identify possible chromatic contributions to spatially induced blackness. Stimuli consisted of a series of monochromatic (400-700 nm; 10-nm steps), 52-arcmin circular test lights surrounded by broadband (x = 0.31, y = 0.37), 63-138-arcmin annuli of fixed retinal illuminance. The stimuli were imaged on the fovea in Maxwellian view as 500-ms flashes with 10-s interstimulus intervals. Observers decreased the intensity of the test center until it was first perceived as completely black. Action spectra determined for two surround levels [2.5 and 3.5 log trolands] had three sensitivity peaks (at approximately 440, 540, and 600 nm), However, when monochromatic surrounds were adjusted to induce blackness in a broadband center, action spectra were unimodal and identical to functions obtained by heterochromatic flicker photometry. Tests of additivity revealed that when blackness is induced by broadband surround into a bichromatic center, there is an additivity failure of the cancellation type. This additivity failure indicates that blackness induction is influenced, in part, by signals from opponent-chromatic pathways. A quantitative model is presented to account for these data. This model assumes that blackness induction is determined by the ratio of responses to the stimulus center and the annulus, and while signals form the annulus are based only on achromatic information, responses from the center are based on both chromatic and achromatic properties of the stimulus.

Contributions of neural pathways to age-related losses in chromatic discrimination.

Chromatic-discrimination thresholds were measured for light mixtures lying along individually determined tritan axes and an axis of constant short-wavelength-sensitive- (S-) cone stimulation for 30 color-normal observers (age range 22-77 years). The stimulus was a foveally viewed 2 degrees, circular bipartite field consisting of a standard and a test light. Heterochromatic flicker photometry was used to equate the retinal illuminance of the stimuli at 120 Td for all observers. All stimuli were presented in Maxwellian view. Age-related losses in chromatic discrimination depended on the level of cone stimulation. At relatively lower levels of S- and long-wavelength-sensitive (L-) cone stimulation, discrimination thresholds were elevated for older relative to younger observers. As the level of simulation increased for these two cone types, thresholds converged, on average, for all observers. Application of a model of chromatic discrimination mediated by an S-cone pathway suggests that there is no significant age-related change in Weber fractions and that age-related losses in chromatic discrimination are due, at least in part, to spontaneous neural noise arising in the pathway and/or neural changes that multiplicatively scale all incident light.

Loci of achromatic points throughout the life span.

Mixtures of monochromatic lights that appear achromatic were measured for 50 normal, trichromatic observers ranging in age from 11 to 78 years. Stimuli were presented to one eye as a 1 degree-diameter, 1-s flash (10-s interstimulus interval) in Maxwellian view. We found the achromatic locus by varying the intensity ratio of each observer's spectral unique blue and unique yellow while maintaining constant overall retinal illuminance. Measurements were made for three levels of retinal illuminance (10, 100, 1000 trolands). Additional verification of the position of the achromatic locus in color space was obtained for 23 subjects with the use of a mixture composed of 600-nm light and its spectral complement. There were no significant changes in the achromatic loci as a function of age. The mean achromatic locus in CIE chromaticity space was x, y = 0.31, 0.31 or u',v' = 0.21, 0.46. These results suggest that partial compensation for age-related changes in visual mechanisms occurs in a way that preserves constancy of the achromatic locus across the life span.

Sites of age-related sensitivity loss in a short-wave cone pathway.

The age-related losses in the sensitivity of a short-wave (S) cone mechanism were determined by measuring the threshold-versus-radiance (t.v.r.) functions for nine older (mean age, 71.0 years) and six younger (mean age, 24.4 years) subjects. Measurements were obtained for a 250-ms, 1.03 degrees-diameter, 440-nm foveally viewed test light presented on 470-nm adapting fields and on a 570-nm auxiliary field. A comparison of t.v.r. functions revealed intensity-dependent sensitivity losses in the older group. By applying a quantitative model of an S-cone pathway to the t.v.r. data obtained from the older subjects, we found that the age-related changes in the ocular media and the receptor sensitivities can account for much but not all of the difference in the sensitivity of an S-cone mechanism demonstrated by eight of our nine older subjects.

Loci of spectral unique hues throughout the life span.

Spectral unique hues (blue, green, and yellow) were determined for 50 observers ranging in age from 13 to 74 years. Each unique hue was measured at three luminance levels (0.5-log-unit steps). There were no significant changes in the spectral locations of red-green equilibrium hues (unique blue and yellow) as a function of luminance level or age. In contrast, significant shifts in unique green loci occurred as a function of both age and luminance. Unique green loci shifted toward shorter wavelengths with age. These results are consistent with the hypothesis that with advancing age there is a parallel decline in the input of all three cone types to the red-green chromatic channel and either a selective decline in short-wave-sensitive cone input to the yellow-blue chromatic channel or a change in the way in which cone signals are combined within the yellow-blue channel.