The influence of cone adaptation upon rod mediated flicker

The influence of annular fields on sensitivity to sinusoidal flicker was assessed in the dark adapted parafoveal retina. Test stimuli were 2 degrees 20' in diameter; annuli had a 2 degrees 20' inner and 7 degrees 30' outer diameter. Rod flicker was studied with a "green" stimulus too dim to influence cones. Selective cone flicker was obtained using red and green flicker in counterphase and yoked together in modulation depth and scotopic illuminance. Results showed the following. (1) Annular stimulation of rods slightly facilitated rod-mediated flicker sensitivity to frequencies less than 10 Hz. In contrast, annular stimulation of cones greatly facilitated rod-mediated sensitivity, particularly for flicker frequencies greater than 7 Hz. We designate this effect, cone-rod interaction. (2) Annular stimulation of cones has a negligible influence upon sensitivity to cone-mediated flicker frequencies less than 15 Hz. In contrast, annular stimulation of rods has a large influence upon sensitivity to cone-mediated flicker, an effect we designate rod-cone interaction. (3) Within limits, both rod-cone and cone-rod interaction increase as the annular illuminance increases and as flicker frequency increases; the limiting frequency and illuminance values, however, are different for the two forms of interaction. Results are compared with prior evidence that rod and cone signals summate to produce an absolute threshold or flicker sensation. We suggest that there are at least three mechanisms for interaction between rod- and cone-related signals.     

Frequency-doubling illusion under scotopic illumination and in peripheral vision

PURPOSE: The authors sought to determine whether frequency-doubling illusion (FDI) could be perceived under scotopic illumination at central and peripheral retinal locations. For comparison, perception of the FDI at the central and peripheral retina under photopic illumination was also evaluated. METHODS: Five subjects matched the apparent spatial frequency of counterphase flickering sinusoidal gratings with stationary sinusoidal gratings presented foveally and out to 20 degrees eccentricity under photopic and scotopic illumination conditions. Two spatial frequencies (0.25 and 0.50 cpd) were used at four temporal frequencies (2, 8, 15, and 25 Hz). Subsequent experiments explored the range of spatial and temporal frequency stimulus conditions under which the scotopic FDI might be observed. RESULTS: Under scotopic illumination conditions, the apparent spatial frequency of eccentrically presented 0.25- and 0.50-cpd flickering gratings gradually increased as a function of flicker frequency and approaches "doubling" at 15 Hz. Under photopic conditions, the apparent spatial frequency of 0.25-cpd flickering at 25 Hz was approximately doubled in all four primary meridians at central and peripheral eccentricities. The final experiment showed that the spatiotemporal range under which the scotopic FDI could be seen was similar to the photopic illumination condition reported earlier. CONCLUSIONS: Scotopic FDI is similar to photopic FDI at the central and the peripheral retina. This suggests that similar mechanisms are responsible for generating the illusion under both photopic and scotopic illumination conditions.     

The temporal properties of the human short-wave photoreceptors and their associated pathways

Flicker modulation sensitivity measurements made on high intensity orange steady backgrounds indicate that signals from short-wavelength sensitive cones (S-cones) have access to two pathways. At low S-cone adaptation levels the frequency response falls quickly with increasing frequency, but at higher adaptation levels it extends to much higher frequencies. At these higher S-cone adaptation levels, the following procedures can selectively expose either a process sensitive to low frequencies or one more sensitive to higher frequencies: (1) at high flicker frequencies, the S-cone signal can be nulled by a long-wavelength sensitive cone (L-cone) signal of suitable amplitude and phase, but at low frequencies a residual flicker persists; the modulation sensitivity for the residual flicker is lowpass in shape with a rapid decline in sensitivity with increasing flicker frequency; (2) sensitivity to flicker in the presence of a 17 Hz S- or L-cone mask is also lowpass with a similarly steep loss of high frequency sensitivity; yet (3) sensitivity to flicker during transient stimulation of the S-cones at 0.5 Hz is comparatively wideband (and slightly bandpass) in shape. The S-cone signal produced by the high frequency process is almost as well-maintained towards high frequencies as M- and L-cone signals. Furthermore, it is capable of participating in flicker photometric nulls with M- and L-cone signals. At low frequencies, however, when the low frequency S-cone signal is also present, satisfactory nulls can not be found. From these and phenomenological considerations, we identify the low and high frequency S-cone processes as S-cone inputs to the chromatic and luminance pathways, respectively. The phase adjustments needed to optimize flicker photometric nulls reveal that the S-cone input to the luminance pathway is actually inverted, but this is demonstrable only at relatively low frequencies: at medium or high frequencies the S-cone influence can be synergistic with that of the other cone types because of a delay in the transmission of S-cone signals.     

Influence of rod adaptation upon cone responses to light offset in humans: I. Results in normal observers.

Dark-adapted rods exert a tonic suppressive influence upon cone-mediated sensitivity to rapid flicker, a phenomenon called suppressive rod-cone interaction (SRCI). However, rod dark adaptation has negligible influence upon cone-mediated thresholds measured with more usual psychophysical procedures. The present study separately examined the influences of rod light and dark adaptation upon cone-mediated sensitivity to transient increases or decreases in illumination using sawtooth flicker with rapid-on (ramp-off) or rapid-off (ramp-on) waveforms. In the parafoveal retina, cones alone were stimulated with flicker by spatially superimposing long- and short-wavelength stimuli presented in counterphase and matched in scotopic illuminance. Several different adaptation procedures were used. For higher (greater than 4 Hz) frequencies, sensitivity of cones to both waveforms is nearly identical under any condition of adaptation; sensitivity decreases as rods progressively dark adapt. A considerably different situation exists for slower frequencies (1-4 Hz). Sensitivity of cones to rapid-off flicker is appreciably greater under light-adapted conditions confirming recent observations by Bowen et al. (1989). But as rods progressively dark adapt, sensitivity of cones to rapid-off waveforms decreases considerably while sensitivity to rapid-on waveforms is much less affected; in the totally dark-adapted eye, sensitivity to both waveforms is identical. These results confirm and extend recent physiological observations in amphibian retina (Frumkes & Wu, 1990) suggesting that SRCI specifically involves responses to transient decreases in illumination.     

Velocity discrimination in scotopic vision

To characterize scotopic motion mechanisms, we examined how variation in average luminance affects the ability to discriminate velocity. Stimuli were drifting horizontal sine-wave gratings (0.25, 1.0 and 2.0 c/deg) viewed through a 2 mm artificial pupil and neutral density filters to produce mean adapting levels from 2.5 to -1.5 log photopic trolands. Drift temporal frequency varied from 0.5 to 36.0 Hz. Grating contrasts were either three or five times direction discrimination threshold contrasts at each adaptation level. Following 30 min adaptation, two drifting gratings were presented sequentially at the fovea. Subjects were asked to indicate which interval contained the faster moving stimulus. The Weber fraction for each base temporal frequency was determined using a staircase method. As previously reported, velocity discrimination performance was most acute at temporal frequencies of about 8.0 Hz and greater than 20.0 Hz (though there are individual differences), and fell off at both higher and lower temporal frequencies under photopic conditions. As adaptation level decreased, discrimination of high temporal frequencies in the central retina became increasingly worse, while discrimination of low temporal frequencies remained largely unaltered. The overall scotopic discrimination performance was best at about 3.0 Hz. These results can be explained by a motion mechanism comprising both low-pass and band-pass temporal filters whose peak and temporal cut-off shifts to lower temporal frequencies under scotopic conditions.     

Psychophysical studies of monkey vision. I. Macaque luminosity and color vision tests

Macaque monkeys and normal human observers were tested on the same apparatus for the presence of a Purkinje shift, and for spectral sensitivity under scotopic and photopic conditions. The flicker fusion point for different lights was used as a measure of visual sensitivity in these tests and the testing procedure was a four-alternative forced choice. The results show that macaque monkeys and normal human observers show a rod-cone break at the same flicker frequency, and are very similar in both relative and absolute scotopic and photopic sensitivity.Macaque monkeys, normal human observers and color-defective human observers were also tested under identical conditions in several measures of color vision.     

High frequency limitations on mach bands

The contrast required to see Mach bands in a trapezoidal grating has been measured under different conditions. At photopic levels, increasing contrast is required as grating frequency increases. Mach bands are absent at all frequencies in scotopic viewing. Their absence at lower frequencies is not due simply to lack of suprathreshold contrast since at low frequencies contrast sensitivity is little affected by dark adaptation.Trapezoids of diminishing ramp width require increasing contrast until for a square edge a contrast of just greater than 1 would be required. This is difficult to achieve in most subjects.The Chevreul illusion behaves quite differently being unaffected by dark adaptation and grating frequency.     

The spatial organization of retinal receptive fields in light and darkness as revealed by the pattern electroretinogram

The spatial selectivity of the electroretinogram in response to pattern onset-offset stimuli was studied in man at several levels of adaptation ranging from scotopic to photopic levels. Under conditions of rod function the peak of the spatial selectivity based on amplitude measurements of the pattern-onset response occurs at a low spatial frequency. With increasing light adaptation a gradual shift of the selectivity to higher spatial frequencies occurs. This change in the character of the response can be explained by the assumption that antagonistic center-surround retinal receptive fields contribute to the response, which are larger under scotopic than under photopic levels of stimulation.     

Using scotopic and photopic flicker to measure lens optical density

Many applications require knowledge of lens absorption. Measuring lens optical density (OD), however, is often difficult and time-consuming. For example, psychophysical measurement typically requires a long period of dark adaptation (e.g. about 40 min) and assessment of absolute scotopic thresholds. In this study, we examined efficient scotopic and photopic methods for measuring lens OD. In Experiment 1, 30 subjects were tested using a Maxwellian-view optical system. Relative scotopic thresholds were obtained after 15 min of dark adaptation using slow-rate (2 Hz) flicker photometry. A 3 degrees test stimulus, presented at 10 degrees nasal, was used that alternated between measuring wavelengths (420 and 460 nm) and a reference field (540 nm). The results showed that the relative scotopic method produces values that are consistent with published lens spectral curves. In Experiment 2, relative photopic measures (i.e. no dark adaptation period) were also obtained at 406 nm in natural view and compared with lens data obtained in Maxwellian view at 407 nm using the absolute scotopic method. The photopic method compared well with the absolute scotopic values obtained on the same subjects. Taken together, the two experiments showed that a relative method can yield valid lens density estimates. Using a relative rather than an absolute method reduces the time needed for dark adaptation and is an easier task for subjects to perform and may therefore be preferable when expeditious measures are desirable.     

Spatial extent of rod-cone and cone-cone interactions for flicker detection

Over a large range of light adaptation levels, sensitivity to 25 Hz flicker improves as the light level of the background increases. Using small background discs and annular surrounds, this effect was shown to be mediated by the surround and not the average luminance of the test region, in agreement with recent reports. The effect is due to two types of lateral interaction: at mesopic light levels (from 0.1 to 1.0 td), cone-mediated flicker resolution is enhanced by the stimulation of surrounding rods; at photopic light levels (above 10 td), flicker sensitivity improves with light stimulation of adjacent cones. The spatial zone, or extent, over which the surround contributes to the flicker threshold was measured. The spatial area over which rods influence the cone 25 Hz flicker threshold is larger than the analogous spatial area of cone influence. In the parafovea, at 5 deg eccentricity, cone flicker sensitivity for a 20' spot is influenced by cones in a 1 deg diameter area centered on the spot; the corresponding area of rod influence is about 3 deg. In the fovea, flicker sensitivity for a 10' spot is influenced by cone stimulation in an area of about 20' diameter. Rods which affect foveal flicker sensitivity appear to occupy an annular zone with about a 2 deg outer diameter and 1 deg inner diameter, centered on the fovea.     

Measuring and modelling the photopic flicker sensitivity of the chicken (Gallus g. domesticus).

The photopic flicker sensitivity of the chicken was determined using an operant conditioning psychophysical technique. The results show both high- and low-frequency fall-off in the sensitivity response, which peaked around 15 Hz. Flicker sensitivity was determined for a range of stimulus luminance levels, and directly compared to human flicker response measured under similar stimulus conditions. At five luminance levels (10, 100, 200, 500 and 1000 cd/m(2)), the overall chicken flicker sensitivity was found to be considerably lower than for humans, except at high frequencies. A greater degree of frequency tuning was also found in the chicken response. The critical flicker fusion values were either similar or slightly higher for chickens compared to humans (40.8, 50.4, 53.3, 58.2 and 57.4 Hz vs 39.2, 54.0, 54.0, 57.4 and 71.5 Hz respectively for humans and chickens for increasing stimulus luminance level). A recently proposed model for flicker sensitivity [Vision Research 39 (1999) 533], which incorporates low- and high-pass temporal filters in cascade, was found to be applicable to the chicken response. From this model, deductions were made concerning mechanisms controlling the transfer of temporal information.     

Photopic 30 Hz flicker ERG as a predictor for rubeosis in central retinal vein occlusion

AIMS: To investigate the predictive value of the cone b-wave implicit time in the photopic compared with the scotopic 30 Hz flicker ERG for rubeosis in central retinal vein occlusion. METHODS: 44 patients with a central retinal vein occlusion were examined with full field electroretinogram (ERG). RESULTS: The average implicit time in the photopic 30 Hz flicker ERG in patients who developed rubeosis (n=15) was 38.3 ms. In the patients who did not develop rubeosis (n=29) it was 31.3. The difference is statistically significant (p=0.0000000004). CONCLUSION: The photopic cone b-wave implicit time in the 30 Hz flicker ERG is a good predictor for rubeosis.     

Frequency dependency of temporal contrast adaptation in normal subjects

The aim of this study was to determine the influence of temporal frequency of temporal contrast adaptation on contrast sensitivity in healthy subjects. Temporal contrast sensitivities (TCS) were measured monocularly in seven healthy subjects with a modified ERG full-field bowl stimulator at eight different test temporal frequencies (9, 15, 20, 25, 31, 37, 44, 51 Hz) using a two-alternative-forced-choice strategy. Before each presentation of the test stimulus, a 100% contrast adapting flicker stimulus was presented (frequencies: 9, 15, 20, 25, 31, 37, 44, 51, 100 Hz). At each adapting frequency, a complete set of TCSs was measured. All temporal contrast sensitivities decreased with increasing temporal frequencies. Adaptation led to a general temporal contrast sensitivity decrease. Largest adaptation effects were seen at an adaptation frequency of 25 Hz. Reduction of contrast sensitivity was significantly larger at 25 Hz adaptation than at 9 Hz adaptation (t-test of paired samples, Bonferroni corrected). The results of this study showed a general TCS decrease with the largest effect at an adaptation frequency of 25 Hz. This finding indicates that the contrast adaptation probably occurred in the magnocellular-pathway. In future clinical studies adaptation effects could be investigated in patients with reduced temporal contrast sensitivity.     

The 52nd ISCEV International Symposium Abstract Issue

Purpose

We have monitored retinal function in patients treated for retinoblastoma (primarily, but not exclusively by intra-arterial chemotherapy infusion) by electroretinography (ERG) recordings for the past 7 years. We here present data from 599 ERG studies of 108 patients, in which a complete ERG protocol including both photopic and scotopic recordings was performed, in justification of our frequent practice of reporting primarily 30-Hz photopic flicker amplitude data.

Methods

Patients referred for treatment of retinoblastoma underwent ERG recordings during examination under anesthesia whenever possible: at baseline and following most treatment sessions. Correlations were calculated for the complete datasets between the four primary amplitude response parameters: photopic single flash b-wave, photopic 30-Hz flicker peak-to-trough, scotopic rod-isolating b-wave, and scotopic maximal flash b-wave.

Results

Using our adaptation of the International Society for Clinical Electrophysiology of Vision-recommended standard ERG protocol, ERG responses of eyes of patients with untreated retinoblastoma or following traditional or intra-arterial treatment for retinoblastoma show very high correlations between 30-Hz flicker amplitude responses and three other standard photopic and scotopic ERG response amplitudes. Reductions in ERG amplitudes seen in these eyes following treatment show no significant difference between retinal dysfunction estimated using rod- or cone-dominated responses.

Conclusion

These observations support the use of photopic response amplitudes (especially in response to 30-Hz flicker) as the primary ERG outcome measure in studies of treated and untreated eyes with retinoblastoma when more complete ERG protocols may be impractical.     

Stimulus chromatic properties affect period doubling in the human cone flicker ERG

Period doubling in the full-field cone flicker electroretinogram (ERG) refers to an alternation in waveform amplitude and/or shape from cycle to cycle, presumably owing to the operation of a nonlinear gain control mechanism. This study examined the influence of stimulus chromatic properties on the characteristics of period doubling in order to better understand the underlying mechanism. ERGs were acquired from 5 visually normal subjects in response to sinusoidally modulated flicker presented at frequencies from 25 to 100 Hz. The test stimuli and the pre-stimulus adaptation were either long wavelength (R), middle wavelength (G), or an equal combination of long and middle wavelengths (Y), all equated for photopic luminance. Fourier analysis was used to obtain the response amplitude at the stimulus frequency F and at a harmonic frequency of 3F/2, which was used as the index of period doubling. The frequency-response function for 3F/2 typically showed two peaks, occurring at approximately 33.3 and 50 Hz. However, the magnitude of period doubling within these frequency regions was dependent on the chromatic properties of both the test stimulus and the pre-stimulus adaptation. Period doubling was generally smallest when an R test was used, even though the stimuli were luminance-equated and the amplitude of F did not differ between the various conditions. The pattern of results indicates that the mechanism that generates period doubling is influenced by chromatic signals from both the test stimulus and the pre-stimulus adaptation, even though the high stimulus frequencies presumably favor the achromatic luminance system.     

Flicker adaptation. II. Effect on the apparent brightness of intermittent lights

It was found that an intermittent light must be made more intense after adaptation to a flickering light in order to appear as bright as it does after adaptation to a steady light of the same time-average retinal illuminance. The largest decreases of brightness gain occurred with adapting stimuli of intermediate frequencies (6–11 Hz). Furthermore, the loss of gain produced by an adapting stimulus of intermediate frequency was highly nonlinear in the sense that it was not constant from one test frequency to the next. Both low- and high-frequency adapting stimuli produced relatively small, but reliable, reductions of gain at all test frequencies. The effects of flicker adaptation on brightness gain are compared with published data on the effects of flicker adaptation on temporal modulation sensitivity, and possible mechanisms of flicker adaptation are discussed.     

Analysis of photoreceptor function and inner retinal activity in juvenile X-linked retinoschisis

Thirteen retinoschisis males with genotyped XLRS1 gene mutations were examined by electroretinogram (ERG) techniques to determine photoreceptor involvement and ON-pathway and OFF-pathway sites of dysfunction. Parameters R(max) and logS determined by fitting the mathematical model of the activation phase of phototransduction to the scotopic and photopic a-wave responses, were not significantly different from normal. However, the XLRS photopic a-wave amplitudes were significantly lower than normal across all intensities, consistent with defective signaling in the OFF pathway. Long flash (150 ms) ON-OFF photopic responses showed reduced b-wave amplitude but normal d-wave amplitude, giving a reduced b/d ratio of <1.32 Hz photopic flicker ERG fundamental frequency responses showed reduced amplitude and delayed phase, consistent with abnormal signaling by both the ON- and OFF-pathway components. These results indicate that the XLRS1 protein appears not to affect photoreceptor function directly for most XLRS males, and that ERG signaling abnormalities occur in both the ON- and OFF-pathway components that originate in the proximal retina.     

Human scotopic spatiotemporal sensitivity: a comparison of psychophysical and electrophysiological data

The aim of the study was to investigate spatiotemporal visual functions under scotopic and photopic conditions in order to acquire human psychophysical and electrophysiological data that are comparable with contrast sensitivities based on single-unit recordings in animal experiments. Static and dynamic contrast sensitivities (CSs) and steady-state visual evoked potentials (VEPs) were measured under photopic and scotopic conditions in healthy volunteers. The results from the CS experiment indicated that the inclusion of temporal modulation and the application of scotopic luminance levels uniformly resulted in a relatively increased sensitivity for low spatial frequencies. Similarly, analysis of the second harmonic component of the VEPs demonstrated a shift from band-pass to low-pass functions. These results suggest that, under scotopic conditions, human visuospatial processing is characteristically predominated by the functional activity of the magnocellular pathways.     

Contrast sensitivity in amblyopia due to stimulus deprivation.

Contrast sensitivity functions for sinusoidal gratings of varying spatial frequency and stimulus duration were determined for both eyes of 2 patients with amblyopia due to early occlusion and lid closure. The amblyopic eyes showed reduced contrast sensitivity over a wide range of spatial frequencies and stimulus durations, and the temporal integration time of the amblyopic eye was increased by comparison with the non-amblyopic eye at high spatial frequencies. When the gratings were flickered at 10 Hz the sensitivity for both flicker and pattern detection was reduced in the amblyopic eye over the entire spatial frequency range. Abnormal flicker perception by the amblyopic eyes was also evident in the reduced photopic luminosity functions of these patients.     

ERG Oscillatory Potentials in Infants

Purpose: Study the scotopic and photopic oscillatory potentials (OPs) of the electroretinogram (ERG) in 10-week old infants. Methods: Term-born 10-week old infants (n = 15) and adults (n = 12) were tested. Full-field ERGs were recorded under scotopic and photopic conditions. The records were filtered (75–300 Hz) to demonstrate the oscillatory wavelets. The amplitudes and implicit times of the infants’ OPs were compared to those in adults and also to amplitudes of the saturated photoreceptor responses. Results: In infants, the mean OP amplitudes are similar in scotopic and photopic conditions and do not vary significantly with OP number. Infants’ OPs are significantly smaller than in adults, with scotopic OPs averaging 19% of that in adults and photopic OPs averaging 47%, whereas the amplitudes of the saturated photoreceptor responses are 43% and 66% of those in adults. Mean interpeak intervals are similar in infants and adults, indicating oscillatory behavior at a frequency of 155 Hz in scotopic conditions and 135 Hz in photopic conditions. Conclusions: In young infants, the OPs are relatively immature compared to the photoreceptor responses, with the immaturity of the scotopic responses being more marked than that of the photopic responses.