When the duration of the noise frame increases, the spectral density below the high-frequency cutoff increases, whereas its cutoff frequency decreases in proportion to the frame’s duration. For a range of short noise frame durations, only the former effect was evident as energy threshold for flicker detection increased in proportion to frame duration and kept the nominal
SNR constant. This means that the masking effect of external, purely temporal noise increased with the duration of the noise frame, mimicking white noise, despite its decreasing bandwidth. For longer noise frame durations, temporal noise did not elevate the detection threshold enough to keep the nominal
SNR constant, because the decreasing cutoff frequency began to curtail even those noise frequencies that interfered with detection. The white part of the noise spectrum thus no longer extended across the whole bandwidth used for detection. This is analogous to our previous results on spatial masking.
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At the cutoff frequency corresponding to the duration of the critical noise frame, the average power spectrum of noise was 41% of its maximum (see
equation 2 ), and at higher frequencies it decreased further in an undulating fashion. When the cutoff frequency of the noise fell below the temporal frequency corresponding to the critical noise frame duration, the spectral density of noise averaged over the frequency range of the detector decreased below
N 0. Hence, the zero-frequency spectral density given by
equation 3 is no longer a valid measure of the effective noise. Noise has become nonwhite within the bandwidth of the detection filter.
The increase of the masking effect of external temporal noise with noise frame duration broadens the use of noise in experimental and clinical research. Both diseases of the visual system and the choice of signal parameters can produce poor sensitivity to temporal contrast, which implies that the contrast of external noise is also attenuated. Because high signal contrast is needed for threshold measurement, the remaining contrast range is not sufficient to produce external noise that is stronger than internal noise. If the spectral density of noise is increased by increasing the duration of the noise frame, however, noise contrast can be kept low, and more of the dynamic range of the display is left for the signal. The critical noise frame duration for the signal used must not be exceeded, however.
The critical duration expressed in terms of a fraction of a cycle increased with flicker frequency. The higher the frequency, the fewer noise frames were needed per cycle to mimic white noise. This differs from spatial vision, for which the critical noise check size expressed as a fraction of spatial cycle is constant, independent of the spatial frequency of the signal with constant bandwidth.
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In the chain of processes
1 2 determining flicker sensitivity, the source of the frequency dependence described herein may be the early physiological transformations shaping neural signals or/and the central signal detector. It would be easy to accommodate the results by ad hoc assumptions about the bandwidths of the matched filter or multiple channels. This is not, however, in agreement with the finding that temporal integration is independent of temporal frequency.
15 16 It is, therefore, more likely that the frequency dependence of critical duration originates from known physiological properties of peripheral processing that modify the neural signals before detection.
In a matched filter model, the detector is assumed to be a replica of the signal. If the bandwidth and center frequency of the neural signal at the level of detection were equal to the external signal, the critical number of noise frames per signal cycle should be constant for our signals with constant bandwidth. The results, however, show that detection of flicker at 1.25 Hz was affected by noise up to approximately 4.2 times the nominal signal frequency. The corresponding factors were approximately 2.3, 1.7, and 1.4 at 2.5, 5, and 10 Hz. This means that the lower the nominal center frequency, the more its bandwidth will spread toward higher temporal frequencies.
Considering the known retinal mechanisms, it seems plausible that the power spectrum of the neural signal could spread to frequencies significantly higher than the nominal signal frequency, especially at low flicker frequencies. There are good grounds for thinking that human achromatic flicker detection is based on signals in M-type retinal ganglion cells.
2 9 17 The nonlinear M
Y cells, such as cat Y cells, give both fundamental and frequency-doubled responses to temporal modulation. The relative amplitudes of these components depend on the parameters of stimulation,
18 19 but the frequency-doubled component is relatively favored by large-field stimuli that extend far into the receptive field surround, as in the present experiments. Moreover, linear responses at the fundamental frequency (whether in M
Y cells or in the larger population
20 of M
x cells) would be comparatively weak for large-field flicker at low frequencies.
21 22 Additional neural modulation above the nominal frequency could come from the convergence of inputs from ON-center and OFF-center cells. We do not know how the brain integrates inputs from different cells and cell types, but a simple assumption would be that detection occurs whenever any type of neural response exceeds a certain
SNR. The relative weight of different response types will automatically vary with flicker frequency, and as signal frequencies approach the temporal high-frequency roll-off of the ganglion cell, higher harmonics will be particularly strongly attenuated, and the fundamental will become dominant. With respect to the nature of the neural signal, it is worth pointing out that the threshold percept of flicker without external noise is quite indeterminate and, specifically, does not correspond to the objective signal frequency.
23
Retinal low-pass filtering can in principle shift the spectral peak of the neural response somewhat below the nominal frequency at high temporal frequencies. This would occur if the flicker frequency is on the falling high-frequency limb of the modulation transfer function (of an M-cell, for example) resulting in such severe filtering at and higher than 20 Hz that the amplitude of the signal (and noise) is relevant mainly at frequencies below the nominal frequency of the signal. In agreement with this, at 20 Hz the critical cutoff frequency of the noise appears to be slightly below the expected minimum value of two noise frames per flicker cycle (i.e., the cutoff frequency of noise equal to the flicker frequency).
Flicker detection has also been modeled
3 4 using a multiple channel approach in which detection is mediated by one broadly tuned low-pass channel and one broadly tuned band-pass channel that includes quite low frequencies but rolls off sharply at higher than 20 Hz. If any subset of our signals were detected using just one of these channels, the critical cutoff frequencies of these signals would correspond to the cutoff of the channel. This could be the case for our signals at 1.25 and 2.5 Hz, because their critical noise cutoff frequencies were almost the same (5.3 and 5.8 Hz) and roughly correspond to the cutoff of the broadly tuned low-pass channel.
4 The increase of critical noise cutoff frequency to 14 Hz when our signal frequency increased to 10 Hz could reflect a combination of the responses from the two channels.
4 Our 20-Hz signal could be detected using only the band-pass channel, because its peak response was at approximately 20 Hz, and at this frequency the response of the low-pass channel had dropped to one tenth of its maximum. The total amount of noise that affects detection should therefore start decreasing as soon as the noise cutoff frequency decreases below the cutoff frequency of the band-pass channel. The critical cutoff frequency of noise therefore should be above the nominal signal frequency, in disagreement with our experimental finding of 16 Hz.
We conclude that the masking effect of external, purely temporal noise can be increased by increasing the noise frame duration up to a critical value. However, the critical duration, in terms of a fraction of flicker cycle, increases with flicker frequency, in contrast to the situation in analogous experiments on spatial masking.
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