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Comparison of the spatial response properties of the human retina and cortex as measured by simultaneously recorded pattern ERGs and VEPs.

Electroretinograms and visual evoked potentials were simultaneously recorded from adult subjects using a checkerboard pattern stimulus reversing at 0.94, 3.75 and 7.5 Hz. Two contrast levels were used: 30 and 85%. The data obtained from the cortex (VEPs) show spatial tuning properties for all temporal frequencies at both contrast levels, with the peak of the amplitude-check size function occurring between 15 and 30 min. Tuning properties were found at the retina but only at the high contrast level and for the faster (3.75 and 7.5 Hz) temporal frequencies. The results demonstrate that spatial tuning is present in the human retina but not under as wide a range of conditions as found at the cortex.

Adult↗

Spatial frequency tuning in the visual evoked potential elicited by sine-wave gratings.

This investigation examined the onset response of the transient visually evoked potential elicited by the appearance-disappearance of sine-wave gratings at various levels of spatial frequency, suprathreshold contrast, and stimulus duration. The response was most easily characterized by two negative-positive complexes that were differentially tuned to spatial frequency. The earlier complex peaked at high spatial frequencies while the later complex peaked at low spatial frequencies. For both complexes, amplitude showed only slight variations across three-octave ranges of contrast and duration. Latency was curvilinearly related to spatial frequency, decreased with increasing contrast, and showed no apparent change as a function of duration.

Adult↗

Responses of neurons in visual cortex (V1 and V2) of the alert macaque to dynamic random-dot stereograms.

A substantial proportion of both simple and complex neurons in the cortex subserving central vision are differentially sensitive to binocular disparity of isolated line patterns (local stereopsis), a sensitivity based on a positional disparity between the neuron's receptive fields in the two eyes. In addition, a subset of cortical neurons, nearly all complex neurons, responds to dynamic random-dot stereograms containing no depth cues other than disparity. These neurons are capable of signaling the correct binocular matches among a multitude of false matches in the stereograms (global stereopsis). The discovery of cyclopean neurons in striate cortex, at early stages of the processing neural network for stereoscopic vision provides a new insight of the basic neural mechanisms underlying binocular depth perception.

Animals↗

The topography of scalp potentials evoked by pattern pulse stimuli.

Pattern pulse stimuli evoke activity in two distinct regions of the human visual cortex in temporal sequence. The earlier responding region (implicit time 93 msec) is localized to a relatively small region near Oz. Its topography is very sensitive to the position of the evoking stimulus in the visual field; it is dominated by a representation of the visual field near the vertical meridian. The later responding cortical region (implicit time 131 msec) is larger than the earlier one. Its medial border touches the lateral border of the earlier responding cortical region and it extends several cm laterally. Its topography is much less sensitive to the position of the evoking stimulus in the visual field. Stimuli presented to the ipsilateral field evoke either weak or no measurable activity in the earlier responding cortical region but often evoke measurable activity near the lateral boundary of the later responding cortical region. These results suggest that the earlier responding cortical region is striate cortex and that the later responding cortical region is composed of visual areas V2, V3, V3A, and V4.

Brain Mapping↗

Visual persistence of stereoscopic stimuli: electric brain activity without perceptual correlate.

Dynamic random-dot stereograms (RDS) were used to study cortical neuronal mechanisms related to visual persistence in nine subjects. Electric brain potential components evoked by stereoscopic stimuli were compared to those evoked by conventional checkerboard stimuli with contrast borders. In a pattern-onset/offset presentation visual persistence thresholds were significantly lower for stereoscopic than for contrast stimuli: with temporally modulated patterns stereoscopic stimuli appeared to visually persist at much shorter interstimulus intervals than contrast stimuli. With stereoscopic stimuli all subjects reported not seeing changes of the RDS pattern, while the corresponding evoked potentials showed components related to the stimulus changes indicating a consistent discrepancy between psychophysical and electrophysiological data. The electrical brain activity was not caused by vergence eye movements elicited by the stereoscopic stimulus. In addition, for dynamic RDS stimuli a significant inverse linear relationship between temporal modulation frequency and evoked potential amplitude was found which was not observed with comparable contrast stimuli.

Adult↗

Electro-physiological investigation of edge-selective mechanisms of human vision.

This study investigates the spatial and temporal characteristics of human visual mechanisms that respond selectively to the polarity of edges. The technique was to record steady-state visual evoked-potentials (VEPs) while visually stimulating with a sawtooth waveform (a series of edges of the same polarity) periodically reversing in contrast (and hence edge-polarity) at a suitable frequency. To ensure that phase-locked VEPs resulted from polarity reversal (rather than local luminance modulation) the stimuli were randomly jittered to a new position between each contrast reversal. The jittered stimulus elicited strong and reliable second-harmonic modulation, usually about one-fifth the amplitude of standard VEPs under similar conditions. The amplitude and extrapolated thresholds of polarity-specific VEPs (relative to standard VEPs) did not vary with eccentricity (up to 10 degrees) or with stimulus orientation. The dependency on spatial frequency was similar to that of standard VEPs, but the polarity-specific VEPs tended to peak at lower temporal frequencies. Perhaps the clearest difference in the two types of VEPs was in the estimated response latency, about 140 msec for the polarity VEPs, compared with 90 msec for standard VEPs.

Contrast Sensitivity↗

The phase of PVEP in Maxwellian view: influence of contrast, spatial and temporal frequency.

The temporal phase of the pattern reversal VEP has been investigated using stimulation with laser interference fringes in Maxwellian view. VEP phase was almost constant as function of spatial frequency (2-30 c/deg, 6.5 r/sec, 51 subjects). The phase function however shows a small phase increase at low spatial frequencies consistent with the existence of multiple temporal mechanisms. Contrast variation yields a smaller phase increase with decreasing contrast than conventional stimulation. The phase is linear as function of reversal rate (2-31 r/sec) for low and high spatial frequencies. The indication of more than one temporal mechanism has also been found when the test field diameter was varied. With decreasing test field size the phase increases at 6.5 r/sec, but decreases at 18 r/sec (for 12 c/deg).

Adult↗

Electrophysiological correlates of texture segregation in the human visual evoked potential.

We investigated whether the visual evoked potential (VEP) reflects cortical processing associated with preattentive texture segregation. On a visual display unit we presented stimuli with various arrangements of oriented line segments that either led to the appearance of a "preattentive" checkerboard or did not. Two presentation modes were used (pattern onset at 1 Hz and rapid pattern change at 4.3 Hz), while luminance (57 cd/m2) and contrast (92%) of the line segments remained constant. VEPs were recorded in 7 human subjects. The VEP was analyzed as a linear combination of putative components, which are evoked by either local pattern, quasi-local orientation contrast or global preattentive structure. In the transient VEP, we found a negativity over the posterior pole at a latency between 161 and 225 msec (FWHM) in the linear combination designed to extract segregation-specific components. Peak amplitude reached 3.1 +/- 0.8 microV (mean +/- SEM) at 199 msec. This negative peak appeared only for textures containing orientation contrast. Steady-state analysis of the rapid presentation also revealed a significant component (P = 0.002) associated with texture segregation. These potentials either represent processing of orientation contrast or global processing of texture segregation. The results suggest that specific surface potentials, differing from cognitive potentials, can be derived which are associated with preattentive processing.

Adult↗

Stereopsis and binocularity in the squirrel monkey.

The squirrel monkey lacks anatomically demonstrable ocular dominance columns, and physiologically it has an ocular dominance distribution in V1 that is very different from that of macaques, with far fewer cells that strongly favor one eye over the other. We tested an alert squirrel monkey for physiological responses to stereoscopic stimuli by measuring evoked potentials in response to cytclopean patterns generated in dynamic random-dot stereograms. The monkey showed evoked responses both to changes in disparity and to shifts between correlation and uncorrelation between the two eyes. This result strongly suggests that the squirrel monkey can detect stereoscopic depth, which in turn casts some doubt on the assumption that ocular dominance columns bear an important relation to stereopsis.

Animals↗

A pupillometric correlate of scotopic visual acuity.

While not easily fit into the classic descriptions of the pupillary light reflex, previous studies reported that changes in the spatial composition of the retinal image can evoke a pupillary response. The present study extends this observation by showing that the pupil constricts in response to scotopic as well as photopic spatial patterns. Moreover, the amplitude of the scotopic response decreases with increasing spatial frequencies suggesting a pupillary spatial acuity of about 3 c/deg. The scotopic pupil acuity is similar to the scotopic perceptual visual acuity measured in the same observers.

Adult↗

Spatial-frequency specific contrast gain and flicker masking of human transient VEP.

We studied the effects of grating contrast and luminance-flicker masking on the early waves of human visually evoked potentials (VEPs) recorded at the onset-offset of sinusoidal gratings of varying spatial frequencies (SFs). At high SFs, the response waveform was simple and VEP was dominated by a negative wave (N110). At low SFs, several positive-negative deflections were recorded, the earliest dominating wave being positive (P90). The amplitude of P90 was saturated at a contrast of about 0.1 and it was attenuated by flicker masking. Masking involved to a lesser extent the waves following P90. It was weaker at the flicker frequency of 5 Hz than at 10 and 20 Hz. No flicker masking was found at SFs higher than 2-4 c/deg. At medium and high SFs, VEPs were obtained at higher contrast levels. No saturation (max contrast tested 0.5) and no flicker masking of N110 were observed. These results suggest that the early VEP components recorded at low and high SFs are related to different types of neuronal activity. Correlation between VEP properties and properties of magnocellular and parvocellular pathways is considered with an emphasis on recent morphological data about the human retina.

Adolescent↗

Visual evoked potentials following abrupt contrast changes.

The timing of visual evoked potential (VEP) amplitude and phase changes following abrupt increases or decreases in contrast was examined. Gratings (1 c/deg) were presented at a low contrast for 8 sec, increased to a higher contrast for 8 sec, and then decreased to the initial lower contrast for another 8 sec. Second harmonic VEP amplitude and phase were recorded continuously and averaged in 1 sec epochs. Both amplitude and phase exhibited delays in reaching a stable level following the contrast change. For amplitude, the length of the delay was dependent on the magnitude and direction of the contrast step and on the spatial frequency of the stimulus. Time constants for the change in amplitude following step increases in contrast ranged from 0.2 sec for a 12% contrast step to 1.34 sec for a 37% contrast step. The timing of phase changes, however, was independent of the size of the contrast increases (tau = 0.7 sec). For step decreases in contrast, both amplitude and phase were relatively independent of the size of the change (tau = approx. 0.9 sec for amplitude and tau = 0.15 sec for phase). Amplitude time constants also increased with increasing spatial frequency (tau = 1.2 sec for 1 c/deg, tau = 1.6 sec for 4 c/deg and tau = 2.3 sec for 8 c/deg); phase time constants, however, did not change as a function of spatial frequency (tau = 0.7 for all spatial frequencies). These findings demonstrate that a unitary process may not always be tapped by signal averaging techniques. Additionally, swept stimulus VEP techniques may produce considerable errors in threshold estimation depending on the stimulus spatial frequency and on the slope and direction of the contrast change.

Contrast Sensitivity↗

Differences between stereopsis, interocular correlation and binocularity.

In normal human subjects, evoked potentials in response to depth reversing two-color dynamic random-dot stereograms disappeared or were greatly reduced at equiluminance, whereas responses to shifts between patterns that were correlated and anticorrelated (for the two eyes) were, for most subjects, actually larger at equiluminance than at non-equiluminance. Responses were only slightly diminished at equiluminance to similar texture-shifting patterns that were identical to the two eyes. These results suggest that a significant fraction of cells with input from both eyes can respond to correlation/anticorrelation shifts, yet are not involved in stereopsis. Also, binocular rivalry may gate the responses of these binocular-nonstereoscopic units.

Depth Perception↗

Linear and nonlinear contributions to step responses in cat retinal ganglion cells.

We measured excitatory and inhibitory step responses of cat retinal ganglion cells to square wave contrast reversal of stationary sinusoidal gratings. In most Y-cells the initial increase in firing rate (early peak) of the excitatory responses was followed by a distinct second increase in firing rate (late peak). Analysis of the spatial frequency and spatial phase dependence of the two peaks indicated that the early peak appears to be produced by the spatially linear center mechanism, while the late peak appears to be produced by the rectifying subunits described by Hochstein and Shapley (1976) Journal of Physiology, London, 262, 237-264, 265-284. These results indicate that the presence of two peaks in ganglion cell step responses is the result of two excitatory inputs with different time courses, and that inhibitory inputs are not required to explain the appearance of these responses.

Action Potentials↗

Development of the temporal properties of visual evoked potentials to luminance and colour contrast in infants.

We have studied the development of the temporal characteristics of the pattern visual evoked potentials (P-VEPs) in response to contrast reversal of patterns of low spatial frequency (0.1 c/deg) of either pure luminance contrast (yellow-black plaid patterns) or pure colour contrast (equiluminant red-green plaid patterns) in 15 infants between 6 and 30 weeks of age. High contrast patterns were modulated temporally either sinusoidally at various temporal frequencies to elicit steady-state responses, or abruptly at a low temporal frequency to elicit transient responses. Analysis of both the transient and steady-state responses suggests the existence of three different mechanisms contributing to the infant and adult P-VEP responses at low, medium and high temporal frequencies. The responses at the three different temporal frequency ranges have different time constants, and develop at different rates. The low frequency response predominates at 8 weeks, where it spans the range 1-6 Hz with an apparent latency of about 230 msec, for both colour and luminance stimulation. This response increases in bandwidth and decreases in latency progressively with age, at a similar rate for luminance and colour contrast, up to 14 weeks. After 14 weeks, the luminance response undergoes major changes, with the emergence of a new response with a shorter latency (about 100 msec) and a peak activity near 10 Hz. This mid-frequency response matures further with age, until it dominates the whole response of the adult P-VEP to luminance contrast. It also makes a contribution to the chromatic response at frequencies above 10 Hz, generating the characteristic double-peaked amplitude response in adults. However, its contribution is very limited below 10 Hz, where the response latency is 140 msec in adults, as it was at 14 weeks of age. A third component is evident at very high temporal frequencies of the luminance response as early as 6 weeks, extending up to 15 Hz in 8-week-olds and up to 25 Hz for older infants. It remains apparent up to 18 weeks, thereafter being swamped by the major mid-frequency response. The apparent latency of response over this frequency range is about 70 msec at all ages. The development of transient P-VEPs paralleled that of the steady-state P-VEPs. At all ages there was an early negative component (N70) at about 70 msec, corresponding to the fast steady-state response at high frequencies for luminance contrast. Before 14 weeks, the luminance and chromatic transient response had the same morphology, with a single major peak of similar latency to the apparent latency of the low temporal frequency response. After this age, the morphology of the luminance response changed, particularly in the first 100 msec, consistent with the emergence of the mid-frequency response. We discuss whether the high-frequency component may represent pre- or early post-synaptic cortical activity, already mature by 8 weeks, and how the different maturation rates of the mid and high-frequency components may reflect different intra-cortical circuitry for colour and luminance.

Adult↗