Brooke E. Schefrin

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.

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.

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.

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.

Spatial summation in human cone mechanisms from 0° to 20° 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°, 4°, 8°, and 20° 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° 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° to 20°, 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.

The area of complete scotopic spatial summation enlarges with age

The maximal area of complete scotopic spatial summation (Riccos 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 Riccos area with age. The average angular subtenses of Riccos 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 Riccos area under scotopic conditions.

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.

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.

Evidence against age-related enlargements of ganglion cell receptive field centers under scotopic conditions

Age-related changes in the neural organization of spatial information are required to account for much of the senescent loss in human scotopic spatial vision, specifically declines in the high spatial frequency cut-off of the contrast sensitivity function and enlargements of the area over which there is complete spatial summation (Riccos area). These results are consistent with hypothesized enlargements of ganglion cell receptive field centers during adulthood. This hypothesis was tested with 50 subjects (19-88 years) by measuring contrast thresholds for two low spatial frequency gratings (0.3 and 1.2 cycles per degree) at a series of scotopic mean illuminance levels. Contrast sensitivity increased with retinal illuminance and then reached a plateau, corresponding to the onset of Weber-like behavior. No age-related change in the light level associated with the onset of Weber-like behavior was found at either spatial frequency. This result is inconsistent with proposed age-related enlargements of ganglion cell receptive field centers under scotopic conditions.

Ricco's Areas for S- and L-Cone Mechanisms Across the Retina

The purposes of this study were to measure areas of complete spatial summation (i.e., Riccos area) for S- and L-cone mechanisms and to evaluate whether the sizes of Riccos area could be explained in terms of either the densities of photoreceptors or ganglion cells. Increment thresholds were measured at the fovea and at 1.5°, 4°, 8°, and 20° in the superior retina using a temporal two-alternative forced-choice procedure. Test stimuli ranging from -0.36 to 4.61 log area (min(2)) were presented on concentric 12.3° adapting and auxiliary fields, which isolated either an S- or L-cone mechanism on the plateau of the respective threshold vs. intensity function. The data indicate that from 0-20° retinal eccentricity, the size of Riccos area is larger for the S-cone mechanism than the L-cone mechanism, increases monotonically for the L-cone mechanism, and, for both cone mechanisms, increases between 8-20° retinal eccentricity. This latter finding suggests that ganglion cell density rather than cone density defines the size of Riccos area in the parafoveal and peripheral retina.