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The application of Laplacian analysis in the recording of half-field pattern-onset evoked potentials.

The Laplacian operator in electroencephalographic measurements consists of a mathematical combination of the responses from a number of electrodes (e.g., five in a crosswise montage). It enhances activity from sources lying underneath the area covered, relative to activity from outside this area. Thus, by appropriate positioning, the contributions of extrastriate and striate sources can be recorded selectively. To quantify the contribution of each hemisphere to half-field onset evoked potentials, the responses in two Laplacian operators, one over each hemisphere, were analyzed and compared to monopolar derivations and a bipolar derivation between the two hemispheres. Both the Laplacian and bipolar analyses were helpful in interpretation of the responses.

Electrodes

Influence of a twofold voluntary hyperventilation on visually evoked cortical potentials and human pupillogram.

We studied the direct and aftereffects of twofold hyperventilation (HV) on pattern reversing VEPs and pupillograms (PGs) of 19 healthy volunteers. The VEP-N80 and P100 latencies increased during HV. Both peak times were maintained for a longer period, up to 20 minutes after HV-2 ended. In addition, the PG-latency time during HV and the PG-construction time during and after HV were increased. The results indicated a temporary delay of neural afferent transmission in the visual system during and after HV. A similar delay of the nervous transmission appeared in the efferent part of the system regulating the pupillary movements after HV ended. The observed changes of the VEP and PG parameters most probably resulted from the hypocapnia cased by HV and its effect on the brain vessels, although other explanations for the changes of the VEP- and PG-parameters may have been possible.

Adult

Spatial frequency characteristics of brisk and sluggish ganglion cells of the cat's retina.

Receptive fields of cat retinal ganglion cells were stimulated by a drifting sinusoidal luminance pattern of fixed (50%) contrast and the amplitude of the fundamental frequency component of response was determined as a function of spatial frequency. Frequency response functions for most cells were unimodal and skewed towards zero frequency when plotted on linear scales. At a fixed retinal location, cells of different classes had different frequency response functions. Heterogeneity within some of the classes could be largely removed by normalizing the axes, thus, revealing a common shape of function for the class. At a fixed retinal location, the maximum response obtained at each spatial frequency was always obtained from a cell of the brisk, rather than sluggish, classes. Spatial frequency resolution was highest for brisk-sustained cells and usually lowest for brisk transient cells.

Animals

Topographic mapping and source localization of the pattern reversal visual evoked magnetic response.

The topography of the visual evoked magnetic response (VEMR) to pattern reversal stimulation was studied in four normal subjects using a single channel BTI magnetometer. VEMRs were recorded from 20 locations over the occipital scalp and the topographic distribution of the most consistent component (P100M) studied. A single dipole in a sphere model was fitted to the data. Topographic maps were similar when recorded two months apart on the same subject to the same stimulus. Half field (HF) stimulation elicited responses from sources on the medial surface of the calcarine fissure mainly in the contralateral hemisphere as predicted by the cruciform model. The full field (FF) responses to large checks were approximately the sum of the HF responses. However, with small checks, FF stimulation appeared to activate a different combination of sources than the two HFs. In addition, HF topography was more consistent between subjects than FF for small check sizes. Topographic studies of the VEMR may help to explain the analogous visual evoked electrical response and will be essential to define optimal recording positions for clinical applications.

Adult

The pattern electroretinogram and visual-evoked potential in glaucoma.

The pattern electroretinogram (PERG) may reflect ganglion cell or inner retinal layer activity. The most sensitive spatial and temporal variables for testing patients with glaucoma have not yet been identified. Fifty-two glaucoma suspects, 51 glaucoma patients, and 28 normal subjects were studied with the PERG and VEP, using three repetition rates and three spatial frequencies. Fast Fourier transforms were calculated at each spatial frequency and reversal rate. An analysis of variance revealed that normals could be differentiated from ocular hypertension and glaucoma patients using the amplitude of the PERG (second and fourth harmonic). Abnormalities in phase of the PERG between groups were also detected. A discriminant analysis of all amplitude and phase data revealed that the phase shift of the response of the second harmonic at 11 alternations/s (15-min checks) and at 5.5 alternations/s (15-min checks) correctly identified 81% of the normal and 75% of the glaucoma patients. The phase shift determinations of the VEP revealed significant abnormalities using 2 and 1/2 standard deviation confidence limits. There was significant overlap in the pattern ERG amplitude and phase shift in all three groups.

Analysis of Variance

Neuronal responsiveness in area 21a of the cat.

Photic responsiveness was studied in cells of area 21a which was identified as a region containing few cells projecting to area 17, and was bounded to two other visual areas (posteromedial lateral suprasylvian area and area 19) providing abundant efferent projection to area 17. Area 21a cells were characterized by strong orientation, but demonstrated poor direction and end-stop selectivity, in contrast to strong direction or end-stop selectivity of cells in another two visual areas.

Animals

Pattern processing and slow waves in visual cortex of visually deprived cats.

The slow-wave response recorded from electrodes within the visual cortex of the awake cat has been shown to reflect the pattern processing activities of the visual cortex. In a cat deprived of pattern vision by monocular lid closure, the response evoked from the non-deprived eye by patterned and diffuse stimuli show differences similar to those found in a normal cat. However, when these same stimuli are presented to the deprived eye, the differences are not present. These slow-wave data confirm previous data from single unit recordings in showing that pattern processing is altered by deprivation.

Animals

Contrast effects of the three primary colors on human visual evoked potentials.

In this study we evaluated in humans the question of whether contrast effects with patterned color stimuli varied in the same way as is known to occur with black-and-white stimuli. Using a counterphasing checkerboard pattern, we evaluated the steady-state visual evoked potential (VEP) in 10 subjects for the response to different contrast levels in each of the 3 primary colors. Overall mean luminance of each color was photometrically equated and kept constant during all trials. The VEP was computer-averaged for 90 consecutive 1 sec epochs of stimulation, and the power at the appropriate frequency was calculated. For each color, the contrast-response curves revealed small power values at low contrast (0.1) and larger power at the 3 high-contrast settings (0.3-0.5). Power varied markedly by color and by subject. The shape of the curves, depending on color and subject, often indicated a saturation of response. A given subject commonly had a physiologically 'preferred' color in that the power with that color was consistently larger. Most subjects had definite subjective color 'preferences,' believing that they perceived the contrast better for one or two colors. However, these impressions were often not validated by the VEP responses to the various colors. These results indicate that white-light VEP responses may not necessarily reflect the response characteristics of specific colors, nor do they necessarily reflect the large inter- and intra-subject differences in color responses noted in this study.

Adult

Human flash-VEP and quantitative EEG are independently affected by acute scopolamine.

Scopolamine in acute intramuscular doses of 0.25-0.75 mg reduced the P2-N3 flash-VEP amplitude and, in the quantitative EEG, the 8.5-12.0 Hz power and total power in 8 healthy young male volunteers. The effects on flash-VEP and EEG total power were dose dependent and were evident 30 min and 90 min respectively after drug administration, regardless of dose. The reduction in 8.5-12.0 Hz power was limited to the 0.50 and 0.75 mg doses. No systematic effects on the pattern-VEP were observed. Possible interferences with flash- or pattern-VEP amplitude of the scopolamine-induced EEG changes were identified and removed by regression analysis and computation of VEP residuals from the regression function. The P2-N3 flash-VEP residuals proved EEG independent and showed relationships with dose and time after drug administration that were superimposable on those of the original data, with comparable significance levels at the drug/placebo and pre/postdrug statistical comparisons. The results indicate that VEP estimates of drug effects which are independent from EEG changes can be identified in human studies and allow some inference on the cholinergic specificity of the systems affecting late flash-VEP components. The statistical approach used in this study is suitable for application in VEP studies when effects of interacting factors are to be expected.

Adult

Contrast sensitivity function and visual acuity of the opossum.

The Modulation Transfer Function (MTF) of the visual system of the opossum, D. marsupialis aurita, was determined using the amplitude of Visually Evoked Cortical Potentials (VECP) as response indicator. Stimuli consisted of a 180 degrees phase reversal of sinusoidally modulated gratings with an average luminance of 2.4 cd/m2. Contrast sensitivity was determined for various spatial frequencies and the MTF was calculated by the least square fit of an exponential function. The average acuity value obtained was 1.25 c/deg. The Fourier transform of the MTF was considered an approximation of the Line Spread Function of the visual system. The lowest value observed was 14 min of arc. The visual acuity observed in the mesopic range was not altered when stimulus intensity was raised to photopic levels.

Animals

Visual acuity in the young infant is highest in a small retinal area.

Visual potentials (VEP) evoked by alternating gratings were recorded daily in one infant between 10 and 13 weeks of age and at 20 weeks of age. The VEP amplitude was measured for a stimulus field of increasing area at low and high spatial frequencies. At low spatial frequencies the VEP amplitude increases with increasing stimulus area, while at high spatial frequencies (near visual acuity) the VEP amplitude remains constant for stimulus field sizes beyond 2 deg. These findings parallel those obtained in the adult, and indicate that visual acuity of the infant is not uniform across the retina but is highest in a small region, as in the adult.

Adult

Ocular refraction and visual contrast sensitivity of the rabbit, determined by the VECP.

The ocular refraction of the rabbit was determined by inserting spherical lenses of between -3 and +4D and measuring the relative VECP amplitude in response to checkboard patterns of 21 min arc. The rabbit's eye was found to be almost emmetropic (+0. 5D ). The contrast sensitivity function of the rabbit's visual system was determined by recording the VECP in response to reversed gratings of different spatial frequencies. The contrast sensitivity was found to be highest at 0.35 c/deg with an upper cut-off frequency at 3 c/deg, corresponding to a grating acuity of 10 min arc.

Animals

Electrophysiological evidence for the existence of coarse and fine disparity mechanisms in human.

Visual sensitivity to stereoscopic disparity changes was measured both psychophysically and by means of evoked potentials. The binocular disparity of a dynamic random-dot stereogram portraying a single flat plane alternated between two values symmetrical about the plane of fixation. The threshold for disparity alternation of the stereoscopic plane was determined at alternation rates between 4 and 12 depth reversals per second (rps). Evoked potential and forced-choice psychophysical estimates of stereoscopic threshold at each reversal frequency agreed, with a mean discrepancy of only +/- 0.1 log units. Evoked potential amplitude was a linear function of log disparity up to about 15 arc min peak to peak disparity. For larger disparities, the evoked potential amplitude versus log disparity function was found to be nonmonotonic with a dip occurring at approximately 26 arc min disparity. Responses to fine disparities of less than 20 arc min lay close to one temporal phase while those evoked by coarse disparities greater than 40 arc min lay near a different phase. The data suggest that disparity processing mechanisms either undergo dynamic changes as disparity increases or that processing shifts between at least two independent mechanisms.

Depth Perception

Representation of edges of variable blur by neuronal responses in the lateral geniculate body and the visual cortex of cats: limits of linear prediction.

We have measured the responses of cells in the cats lateral geniculate body and the visual cortex to edges which were blurred to various degrees (cosinusoidal blur). For the same cells also the responses were determined to sinusoidal gratings of various fundamental frequency and to slits of various blur and width. All stimuli were moved across the receptive fields at various speeds. The responses of most cells increased with increasing edge sharpness, but usually reached a maximum at a blur corresponding to a high frequency cutoff at 0.6-1.2 c/deg. The responses to the sharpest edges were usually smaller than those to a blurred edge (up to -50% in individual cells and -15% in the average). After normalization, the responses predicted from the spatial frequency tuning curves and the Fourier transform of the edge stimuli corresponded well to the measured blur functions up to the maximum of the edge response which varied considerably between cells, however. At edge sharpness beyond that maximum, the predicted curves rose up to edge sharpness with high frequency cutoff 1.6-1.8 times above that which produced the experimental neuronal response maximum. On the other hand, responses could increase with edge sharpening in spatial frequency regions, in which no or only small responses were seen with sinusoidal gratings (e.g. at lower spatial frequencies in "band pass neurons"). Geniculate X- and cortical simple cells as well as those geniculate Y-cells which showed phase locked grating responses behaved similarly in all respects. We concluded that edge sharpness is not represented by response amplitude of individual neurons but by the spatial distribution of excitatory peaks across the representation of the retinotopic cortical map. Our findings further indicate that spatial models of receptive fields assuming linear signal summation have only a limited value for predicting edge sharpness.

Animals

Responses of visual cortical neurons to curved stimuli and chevrons.

Single cells were recorded in area 17 of anaesthetized and paralyzed cats and their responses to curved stimuli and chevrons compared. Striate cells exhibited three different response patterns. A first group responded optimally to a straight line (i.e. zero curvature) and responded similarly to chevrons and to curved lines. A second group responded to all curvatures and was broadly tuned for the straight line when tested with chevrons. A third group responded only to large curvatures, many (2/3) to both signs of curvature and a number (1/3) to only one sign. Cells in this group responded differently to chevrons and curved lines. Cells in these three classes differed both in length-response curve and in width of orientation tuning. Laminar analysis revealed that the three classes are distributed differently across cortical layers. These data shed new light on the finding of Malpeli and coworkers that orientation is extracted at least twice in a cortical column.

Animals

The two-dimensional spatial structure of nonlinear subunits in the receptive fields of complex cells.

We have estimated the second-order response properties of complex cells in two spatial dimensions by cross-correlating their spike trains with a binary approximation of a Gaussian white noise stimulus ensemble. Wiener-like kernels were computed and generally consisted of two or three parallel, elongated subregions alternating between augmented and suppressed response. These subunits were scattered across the receptive fields of complex cells and their axes of elongation agreed with the optimal orientation determined with drifting gratings.

Animals

The visual acuity of the lynx.

Visual evoked potentials were recorded from the occipital scalp of two anaesthetized Lynx (Lynx europea) in response to alternating gratings of various spatial frequencies and contrasts. The visual acuity of the Lynx was found to be around 5-6 c/deg, i.e. very close to the visual acuity of the cat and by far inferior to human acuity.

Animals