This essay advances a version of the flicker of freedom defense of the Principle of Alternative Possibilities (PAP) and shows that it is invulnerable to the major objections facing other versions of this defense. Proponents of the flicker defense argue that Frankfurt-style cases fail to undermine PAP because agents in these cases continue to possess alternative possibilities. Critics of the flicker strategy contend that the alternatives that remain open to agents in these cases are unable to rebuff Frankfurt-style attack on the grounds that they are insufficiently robust (that is, morally significant in a way that could ground ascriptions of moral responsibility). Once we see that omissions are capable of constituting robust alternatives, even when they are not intentional, it becomes clear that agents in these cases do indeed possess robust alternative possibilities—alternatives that are ineliminable from cases of this sort. The upshot is that Frankfurt-style cases are theoretically incapable of providing us with good grounds for rejecting PAP.

This study defined the amplitude and phase characteristics of rod- and cone-isolated flicker electroretinograms (ERGs) and determined how these responses summate to generate the nonreceptor-specific ERG. Full-field ERGs were obtained from six normally sighted subjects (age 26 to 44 years) using a four-primary LED-based photostimulator and standard recording techniques. The four primaries were either modulated sinusoidally in phase to achieve simultaneous rod and cone activation (ERGR+C; nonreceptor-specific) or in different phases to achieve rod-isolated (ERGR) and cone-isolated (ERGC) responses by means of triple silent substitution. ERGs were measured at two mean luminance levels (2.4 and 24 cd/m2), two contrasts (20 and 40%), and four temporal frequencies (2–15 Hz). Fundamental amplitude and phase for each condition were derived by Fourier analysis. Response amplitude and phase depended on the stimulus conditions (frequency, mean luminance, and contrast), however, for all conditions: 1) response phase decreased monotonically as stimulus frequency increased; 2) response amplitude tended to decrease monotonically as stimulus frequency increased, with the exception of the 24 cd/m2, 40% contrast ERGR+C that was sharply V-shaped; 3) ERGR phase was delayed (32 to 210 deg) relative to the ERGC phase; 4) ERGR amplitude was typically equal to or lower than the ERGC amplitude, with the exception of the 2.4 cd/m2, 40% contrast condition; and 5) the pattern of ERGR+C responses could be accounted for by a vector summation model of the rod and cone pathway signals. The results show that the ERGR+C amplitude and phase can be predicted from ERGR and ERGC amplitude and phase. For conditions that elicit ERGR and ERGC responses that have approximately equal amplitude and opposite phase, there is strong destructive interference between the rod and cone responses that attenuates the ERGR+C. Conditions that elicit equal amplitude and opposite phase rod and cone responses may be particularly useful for evaluating rod–cone interactions.

At certain temporal frequencies, the human cone flicker electroretinogram (ERG) contains multiple additional responses following the termination of a flicker train. The purpose of this study was to determine whether these poststimulus responses are a continuing response to the terminated flicker train or represent the oscillation of a resonant system. ERGs were recorded from 10 visually normal adults in response to full-field sinusoidally modulated flicker trains presented against a short-wavelength rod-saturating adapting field. The amplitude and timing properties of the poststimulus responses were evaluated within the context of a model of a second-order resonant system. At stimulus frequencies between 41.7 and 71.4 Hz, the majority of subjects showed at least three additional ERG responses following the termination of the flicker train. The interval between the poststimulus responses was approximately constant across stimulus frequency, with a mean of 14.4 ms, corresponding to a frequency of 69.4 Hz. The amplitude and timing characteristics of the poststimulus ERG responses were well described by an underdamped second-order system with a resonance frequency of 70.3 Hz. The observed poststimulus ERG responses may represent resonant oscillations of retinal ON bipolar cells, as has been proposed for electrophysiological recordings of poststimulus responses from retinal ganglion cells. However, further investigation is required to determine the types of retinal neurons involved in the generation of the poststimulus responses of the human flicker ERG.

Visually evoked field potentials in human subjects and single-cell responses from retinal ganglion cells in the macaque monkey were compared in closely similar stimulus situations. The classical heterochromatic flicker photometry (HFP) technique was used to measure spectral sensitivity in man, both psychophysically and by recording the 40-Hz response, and to measure the spectral sensitivity of magnocellular (MC-) pathway cells of the macaque. The three measures gave closely similar spectral-sensitivity curves. Close agreement between the three measures was also found when the variable-modulation HFP technique was used to measure spectral sensitivity. When the relative phase between red and green lights was varied, the point of minimum subjective flicker for human observers was close to a sharp minimum found in the amplitude of the 40-Hz response in human and was also close to a minimum in the response of MC-pathway neurons in the monkey. The human 40-Hz response saturated at between 10 and 30% modulation depth, and so did the response of MC-pathway cells in the monkey.

The 16-Hz response in human showed none of the above correlations with MC-pathway properties. On the other hand, parvocellular (PC-) pathway cells responded vigorously to constant-luminance, chromatic modulation, at frequencies higher than can be detected by human observers. The human 16-Hz response also was strong in that stimulus situation. In addition, the response of PC-pathway cells on increasing modulation depth showed little saturation, and this behaviour was paralleled by the human 16-Hz response.

We conclude that the properties of MC-pathway neurons in macaque are closely similar to the properties of the human 40-Hz response in the respects just described. We suggest that the 40-Hz response may offer a means of objectively isolating and investigating the contribution of the magnocellular stream to cortical activity in human. In contrast, the properties of PC-pathway neurons in macaque are quite different from the properties of the human 40-Hz response, and in several respects resemble the properties of the human 16-Hz response.

We examined the responsivity, orientation selectivity, and direction selectivity of a sample of neurons in cortical area V1 of the macaque using visual stimuli consisting of drifting oriented contours defined by each of two very different figural cues: luminance contrast and temporal texture. Comparisons of orientation and direction tuning elicited by the different cues were made in order to test the hypothesis that the neuronal representations of these parameters are form-cue invariant. The majority of the sampled cells responded to both stimulus types, although responses to temporal texture stimuli were generally weaker than those elicited by luminance-defined stimuli. Of those units exhibiting orientation selectivity when tested with the luminance-defined stimuli, more than half were also selective for the orientation of the temporal texture stimuli. There was close correspondence between the preferred orientations and tuning bandwidths revealed with the two stimulus types. Of those units exhibiting directional selectivity when tested with the luminance-defined stimuli, about two-thirds were also selective for the direction of the temporal texture stimuli. There was close correspondence between the preferred directions revealed with the two stimulus types, although bidirectional responses were somewhat more common when temporal texture stimuli were used. These results indicate that many V1 neurons encode orientation and direction of motion of retinal image features in a manner that is largely independent of whether the feature is defined by luminance or temporal texture contrast. These neurons may contribute to perceptual phenomena in which figural cue identity is disregarded.

Current approaches to the problem of equating different colors for luminance (chromatic isoluminance) rely upon human reports of perceptual events that are reduced at some luminance ratio. In this report, a technique is described that evokes a vivid percept of motion of a textured pattern only at isoluminance. Furthermore, in both humans and monkeys, the moving stimulus produces a striking optokinetic response in the same direction as the perceived motion. If used in this manner, the technique can provide an estimate of chromatic isoluminance in a variety of species and be used to corroborate a human subjects's perceptual judgement.

Ibotenic-acid lesions of the magnocelluar portion of the macaque lateral geniculate nucleus were used to examine the role of the M-cell pathway in spatio-temporal contrast sensitivity. A lesion was place in layer 1 of the lateral geniculate of each of two monkeys. Physiological mapping in one animal demonstrated that the visual-field locus of the lesion was on the horizontal meridian, approximately 6 deg in the temporal field. Visual thresholds were tested monocularly in the contralateral eye, and fixation locus was monitored with a scleral search coil to control the retinal location of the test target.

Three threshold measures were clearly disrupted by the magnocellular lesions. Contrast sensitivity for a 1 cycle/deg grating that drifted at 10 Hz was reduced from about twofold greater than, to about the same as, that for 10-Hz counterphase modulated gratings. Sensitivity for a very low spatial frequency (Gaussian blob), 10-Hz flickering stimulus was reduced so severely that no threshold could be measured. In addition, flicker resolution was greatly reduced at lower modulation depths (0.22), but not at higher depths (1.0). Two of the measured thresholds were unaffected by the lesions. Contrast sensitivity for 2 cycle/deg stationary gratings remained intact, and little or no effect on sensitivity was found for 1 cycle/deg, 10-Hz counterphase modulated gratings.

Together, these results suggest that the magnocellular pathway makes little contribution to visual sensitivity at low to moderate temporal frequencies. On the other hand, some contribution to detection sensitivity is evident at lower spatial and high temporal frequencies, especially for drifting stimuli. It appears that a major role of the magnocellular pathway may be to provide input to cortical mechanisms sensitive to rapid visual motion.

Difference-of-Gaussians (DOG) models for the receptive fields of retinal ganglion cells accurately predict linear responses to both periodic stimuli (typically moving sinusoidal gratings) and aperiodic stimuli (typically circular fields presented as square-wave pulses). While the relation of spatial organization to retinal anatomy has received considerable attention, temporal characteristics have been only loosely connected to retinal physiology. Here we integrate realistic photoreceptor response waveforms into the DOG model to clarify how far a single set of physiological parameters predict temporal aspects of linear responses to both periodic and aperiodic stimuli. Traditional filter-cascade models provide a useful first-order approximation of the single-photon response in photoreceptors. The absolute time scale of these, plus a time for retinal transmission, here construed as a fixed delay, are obtained from flash/step data. Using these values, we find that the DOG model predicts the main features of both the amplitude and phase response of linear cat ganglion cells to sinusoidal flicker. Where the simplest model formulation fails, it serves to reveal additional mechanisms. Unforeseen facts are the attenuation of low temporal frequencies even in pure center-type responses, and the phase advance of the response relative to the stimulus at low frequencies. Neither can be explained by any experimentally documented cone response waveform, but both would be explained by signal differentiation, e.g. in the retinal transmission pathway, as demonstrated at least in turtle retina.

In the human rod visual system, self-cancellation of flicker signals is observed at high rod intensity levels near 15 Hz, both perceptually and in the electroretinogram (ERG). This and other evidence suggests that two rod signals are transmitted through the human retina with different speeds of transmission. Here we report a series of flicker ERG recordings from a normal observer and an observer who lacks cone vision. From these results, we propose a quantitative model of the two rod signals, which assumes (1) that the amplitude of the slow signal grows linearly with log intensity but then saturates at ~1 scot, td; (2) that the amplitude of the fast signal grows linearly with intensity; (3) that there is a difference in time delay of ~33 ms between two rod signals of the same polarity (or of ~67 ms if the signals are of inverted polarity); and (4) that the time delay of both signals declines linearly with log intensity (by ~10 ms per log scot. td). These simple assumptions provide a remarkably good account of the experimental data. Our results and model are relevant to current anatomical theories of the mammalian rod visual system. We speculate that the slower signal in the human ERG may reflect the transmission of the rod response via the rod bipolars and the An amacrine cells, while the faster signal may reflect its transmission via the rod-cone gap junctions and the cone bipolars. There are, however, several objections to this simple correspondence.

Behavioral and electrophysiological methods were used to measure sensitivity to flickering lights in a dichromatic species, the California ground squirrel (Spermophilus beecheyi). Discrimination tests were used to determine spectral sensitivity at stimulus frequencies from 5–50 Hz and increment threshold spectral sensitivity. The contributions of retinal mechanisms to these capacities were assessed by recording the responses of optic nerve fibers to temporally modulated monochromatic lights. In the ground squirrel, as in the human, the shape of the spectral-sensitivity function depends on the temporal frequency of the stimulus. Results from single-unit recording show that all of the classes of optic nerve fibers in the ground squirrel are highly phase-locked to the stimulus for modulation rates as high as 50 Hz. Neither the responses of photoreceptors nor any class of optic nerve fiber can singly account for the behavioral results. The electrophysiological results are also counter to models which propose that temporally dependent changes in the spectral sensitivity of spectrally opponent fibers account for the behavior. The temporal resolution of the optic nerve fibers exceeds that of the behaving animal suggesting that retinal mechanisms do not limit behavioral temporal resolution.

Omitting a stimulus from a train of repetitive stimuli, by either interrupting or terminating the train, can elicit an electrophysiological response that occurs at the time appropriate for the omitted stimulus. This study investigated whether such an omitted stimulus response (OSR) is present in the flicker electroretinogram (ERG) of the human cone system. ERGs were recorded from 11 visually normal subjects in response to full-field sinusoidal flicker trains presented against a rod-desensitizing adapting field at frequencies ranging from 12.5 to 100 Hz. Recordings were synchronized with the onset of the stimulus trains, and the amplitude and relative delay of any additional ERG responses following the offset of the flicker train were analyzed. At stimulus frequencies below 35 Hz, the number of ERG responses always equaled the number of stimulus cycles. However, over the frequency range of 38.5 to 100 Hz, the ERG contained an extra response following flicker train offset. At stimulus frequencies from 38.5 to 62.5 Hz, there was a constant delay between the peak of the extra ERG response and the time at which the next stimulus would have occurred had the flicker train continued. This constant delay is characteristic of an OSR. In addition, an extra ERG response was apparent at these same stimulus frequencies if the flicker train was interrupted by omitting stimulus cycles from the middle of the train. The pattern of ERG findings is consistent with a recently proposed model of the OSR that attributes the phenomenon to a resonant oscillation in retinal bipolar cells.

Electroretinogram (ERG) responses of the cone system to a flickering stimulus can exhibit a cyclic variation in amplitude. This phenomenon of synchronous period doubling has been attributed to a nonlinear feedback mechanism within the retina that alters response gain. The aim of the present study was to investigate intersubject variability in period doubling in the ERG of the human cone system, and to assess the implications of this variability for signal processing within the retina. Period doubling was examined in a group of 12 visually normal subjects, using sinusoidal full-field flicker and harmonic analysis of the ERG waveforms. For all subjects, the ERG responses to 32-Hz flicker (a frequency commonly used clinically) were characterized by a harmonic component at the stimulus frequency and at higher harmonics that were integral multiples of the stimulus frequency, as expected. In addition, six of the subjects showed period doubling at 32 Hz, characterized by harmonic components at integer multiples of a frequency that was half the stimulus frequency (the subharmonic). However, the subharmonic itself did not exceed the noise level. These findings suggest that the subharmonic is generated prior to or at the site that produces the nonlinear higher harmonics of the ERG response, and that a subsequent band-pass filter attenuates this subharmonic. Examination of harmonic components of the subjects' ERG waveforms at other stimulus frequencies, as well as a cycle-by-cycle analysis of the ERG waveforms, suggested that individual differences in period doubling may be due to intersubject variation in the strength of the hypothesized feedback signal and/or the time constant of its decay.

The retinal dopaminergic system is a global regulator of retinal function. Apart from the fact that the rates of dopamine synthesis and release are increased by increasing illumination, the visual image parameters that influence dopaminergic function are mostly unknown. Roles for spatial and temporal frequency and image contrast are suggested by the effects of form-deprivation with a diffusing goggle. Form-deprivation reduces the rates of dopamine synthesis and release, and induces myopia, which is prevented by dopamine agonists. Our purpose here was to identify visual stimulus parameters that activate dopaminergic amacrine cells and elicit dopamine release. White Leghorn cockerels 4–7 days old were exposed to 2 h of form-deprivation, reduced light intensity, or stimuli of varied temporal or spatial frequency. Activation of dopaminergic neurons, labeled for tyrosine hydroxylase (TH), was assessed with immunocytochemistry for c-Fos, and dopamine release was measured by HPLC analysis of dopamine metabolite accumulation in the vitreous body. Form-deprivation did not reduce TH+ cell activation or vitreal dopamine metabolite accumulation any more than did neutral-density filters of approximately equal transmittance. TH+ cell activation and vitreal metabolite accumulation were not affected significantly by exposure to 2, 5, 10, 15, or 20 Hz stroboscopic stimulation on a dark background, or by sine-wave gratings of 0.089, 0.44, 0.89, 1.04, or 3.13 cycles/deg compared to a uniform gray target of equal mean luminance. These data indicate that the retinal dopaminergic system does not respond readily to short-term changes in visual stimulus parameters, other than light intensity, under the conditions of these experiments.

When a temporally fluctuating background is rapidly modulated (e.g. 30 Hz), the threshold variation of a superimposed flash (the probe) is approximately sinusoidal and in phase with the stimulus. But, with low rates of sinusoidal modulation (e.g. 1 Hz), the threshold variation is distinctly nonsinusoidal in shape. The bases of these aspects of the data, as well as an unmodulated, dc, threshold elevation, are poorly understood. Here 30-Hz and 1-Hz conditions are simulated using a new model of light adaptation (Wilson, 1997). By assuming that the OFF pathway is twice as sensitive as the ON pathway, the model correctly captured the key aspects of both conditions. The results suggest that the 1-Hz data are mediated by a mixture of ON and OFF pathways while the 30-Hz data are largely mediated by the OFF pathway. The probe thresholds on the 30-Hz background appear approximately sinusoidal and approximately in phase with the background stimulus. A number of factors contribute to this deceptively simple observation.