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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

Influence of spatial frequency on tuning and bias for orientation and direction in the cat's striate cortex.

Directionality, orientation and spatial frequency tuning were determined for 108 neurones recorded extracellularly from the striate cortex of anaesthetized cats. Significant sharpening of orientation selectivity with increasing spatial frequency was seen in all simple neurones and the overwhelming majority of complex neurones. Orientation selectivity sharpened in 90 and broadened in only 10 of 100 fully characterized neurones. At least four distinct classes of neurone could be characterized on the basis of their directionality at optimal spatial frequency, and the presence or absence of changes in directionality over a range of spatial frequencies: in two classes, directionality was spatial-frequency dependent; in the remaining two it was invariant. With two exceptions Type A neurones (23 cells) were direction-selective; they were narrowly tuned for orientation and spatial frequency, and their directionality was invariant with spatial-frequency. The majority of neurones (52 cells) were Type B, most of which were direction-biased; their bias for direction varied systematically with spatial frequency. Type C were direction-biased and spatial-frequency selective (5 cells), but showed a clear reversal of bias with change in spatial frequency. Type D, a subset of direction-biased cells, were bidirectional and spatial-frequency invariant (8 cells), with comparable response strengths to motion in two opposing directions at all spatial frequencies. These response types crossed traditional boundaries between categories of simple and complex neurones, assigned on the basis of spatial summation, presence or absence of end-inhibition, and receptive field size.

Animals

Classifying simple and complex cells on the basis of response modulation.

Hubel and Wiesel (1962; Journal of Physiology, London, 160, 106-154) introduced the classification of cortical neurons as simple and complex on the basis of four tests of their receptive field structure. These tests are partly subjective and no one of them unequivocally places neurons into distinct classes. A simple, objective classification criterion based on the form of the response to drifting sinusoidal gratings has been used by several laboratories, although it has been criticized by others. We review published and unpublished evidence which indicates that this simple and objective criterion reliability divides neurons of the striate cortex in both cats and monkeys into two groups that correspond closely to the classically-described simple and complex classes.

Animals

A programmable digital video pattern generator controlled by microprocessor for basic research and clinical applications.

A programmable digital video pattern generator controlled by a microprocessor is described as a part of any commercial or specific visual evoked potential analysis system. The generator handles all kinds of patterns (uniform, orthogonal and radial) with all known presentation modes (onset, offset and reversal) and resolution 640 x 200 square pixels. The frame rate of the generator is adjusted to be compatible to either low cost 50 Hz video monitor or expensive ones of higher frame rate. The operation parameters defined by software are: pattern selection, trigger mode, field selection and stimulation rate. The development of software applications for special research demands is desirable because of the control of the generator by a microprocessor.

Computer Graphics

Texture segregation is processed by primary visual cortex in man and monkey. Evidence from VEP experiments.

We investigated whether the process of texture segregation can be allocated to a specific visual cortical area. We designed a stimulus to reveal the presence of a mechanism, which is specifically sensitive to a checkerboard, that is solely defined by textures segregating due to orientation differences of the constituting line segments. We recorded evoked potentials to this stimulus in man and awake monkey. A difference component, signalling texture segregation sensitivity, could be recorded from both types of subjects. Its presence depended on the spatial extent of the textures, in a manner correlating with the perceptibility of the checkerboard. This difference response could be localized in primary visual cortex by means of equivalent dipole estimations.

Animals

Orientation selectivity of 3-month-old infants.

A modification of the visual evoked potential (VEP) technique, first employed by Braddick, Wattam-Bell and Atkinson [(1986) Nature, London, 320, 617-619] was used to estimate the orientation selectivity of 3-month-old infants. The orientation-selective VEP was recorded in response to various changes in the orientation of a square-wave grating. The magnitude (the square root of the power of the FFT) at the frequency of orientation change was assumed to represent the response to the change in grating orientation. Orientation sensitivity was then estimated by plotting the magnitude of the FFT at the frequency of orientation change as a function of the log of the orientation change in degrees. For each individual, the data were fit by nonlinear regression and threshold was defined as the largest orientation angle for which the magnitude of the FFT was zero. The results suggest that the orientation selectivity of 3-month-old infants (1.33 deg) is similar to that of adults (1.13 deg) tested with the same stimulus parameters (1 c/deg, 9 Hz). However, when adults are tested with stimulus parameters selected to optimize their VEP response (4 c/deg, 18 Hz) instead of those which optimize the infant's response, the orientation discrimination of adults improves by a factor of 2 (0.53 deg). The results obtained from adults under optimum stimulus conditions (4 c/deg, 18 Hz) approach the estimates reported in the literature for static stimuli.

Adult

Cortical neurons: isolation of contrast gain control.

The selectivity of cortical neurons remains invariant with contrast, even though the contrast-response function saturates. Both the invariance and the saturation might be due to a contrast-gain control mechanism. To test this hypothesis, a drifting grafting was used to measure the contrast-response function, while a counterphase grating was simultaneously presented at the null position of the receptive field (where it evokes no response at any contrast). When the contrast of the counterphase grating increased, the contrast-response function shifted primarily to the right. This result is consistent with the hypothesis that there is a fast-acting gain-control mechanism which effectively scales the input contrast by the average local contrast.

Adaptation, Ocular

Rat and human visual-evoked potentials recorded under comparable conditions: a preliminary analysis to address the issue of predicting human neurotoxic effects from rat data.

Pattern-onset visual-evoked potentials (VEPs) were recorded from rats and humans in order to perform cross-species comparison of neuronal functional properties reflected by the early VEP components. The spatial frequency of a sinusoidal test grating was varied in Experiment 1. For both species, amplitude of the first positive VEP component was larger at low spatial frequency and decreased as spatial frequency increased. The immediately succeeding negative component was small at low spatial frequency and was of maximal amplitude at moderate spatial frequency. The effects of stationary pattern adaptation on these components were investigated in Experiment 2. Subjects viewed either a blank field or the test grating prior to recording VEPs. For both species, adaptation had no effect on the positive component but strongly attenuated the negative component. Experiment 3, in which only humans were tested, indicated that the negative component was of cortical origin. Only cortical neurons are known to be orientation selective, and the effect of adaptation diminished as the orientation difference between the adaptation and test gratings increased. These results suggest that the early positive and negative components arise from parallel visual pathways, and that the rat components may reflect visual processes qualitatively similar to those of humans.

Adaptation, Physiological

Asymmetries in ON and OFF visual pathways of humans revealed using contrast-evoked cortical potentials.

Positive- and negative-contrast stimuli yield the perceptions of brightness and darkness, respectively, and are processed separately by ON and OFF neural pathways. The properties of these morphologically and pharmacologically distinct subsystems were measured in humans by recording visual evoked potentials (VEPs). These electrical responses from the visual cortex were elicited by novel positive- and negative-contrast stimuli, designed to emphasize, selectively, contributions from ON and OFF pathways. Results revealed differential processing of the two types of contrast information, suggesting asymmetries in ON and OFF subsystems; OFF subsystems have finer spatial tuning and greater contrast gain than ON subsystems. These VEPs may be useful in diagnosing neurological disorders that involve primarily one subsystem.

Adult

Electroretinograms (ERGs) and visual-evoked potentials (VEPs) elicited by pattern displacement.

The relation between the amplitude of visual responses to a checkerboard stimulus and the degree of lateral displacement of the checks was examined across different check sizes with simultaneously recorded electroretinograms (ERGs) and visual-evoked potentials (VEPs). The amplitudes of both the b-wave and the after-potential of the ERG increase linearly with pattern displacement. However, the major components of the VEP (N70 and P100) were smaller than expected from linearity for both small checks with small displacements (thresholding) and for large checks with large displacements (saturation). These results suggest that the ERG is proportional to the number of receptors stimulated, but the VEP reflects neural processes influenced by the spatial structure of the stimulus.

Adult

Cortical potentials evoked by short wavelength patterned light.

Pattern-evoked cortical potentials (PECPs) were recorded in response to the onset of blue and black, square wave gratings superimposed on a bright orange background. Several lines of evidence, including spectral sensitivity and the absence of response in a tritanopic subject, indicate that the measured cortical potentials reflect input from S cones. Spatial and temporal tuning of this response are comparable to psychophysical measures of the S-pathway. In agreement with previous studies of achromatic and chromatic spatial processing, a comparison of the response to patterns of different complexity indicates that spatial processing in the S-pathway can be described by linear systems analysis. An oblique effect for the S-pathway is demonstrated. Additional findings which bear on the nature of postreceptoral processing in the S-pathway are considered. Included are measurements from a patient with diabetic retinopathy. This study underscores the potential importance of the S-pathway for spatial information processing.

Adult

Multiple sources of P3b associated with different types of information.

This experiment investigated how the P3a, P3b, and Slow Wave components of the event-related brain potential (ERP) respond to manipulations of the nature, timing, and extent of information delivery. There were two experiments in which the total amount of task information was distributed between pairs of successive stimuli (S1 and S2) within each trail. The task was to predict the relation between S1 and S2. In Experiment 1, the S1 could resolve no, partial, or all uncertainty with respect to the prediction outcome (correct or incorrect). Each S1 delivered three types of information: 1) outcome information--which resolved the subjects' uncertainty about the correctness of their prediction; 2) procedural information--which resolved uncertainty about how much outcome information would be delivered by S1; and 3) memory information--the identity of S1, which had to be stored for subsequent comparison with S2. In Experiment 2, the activity of these components was contrasted in two conditions in which the S1 delivered either memory information alone or both memory and procedural information. P3a and Slow Wave were sensitive only to outcome information. P3b was sensitive to all three types of information, and its scalp topography varied as a function of the type of information. The topographic variations indicate that P3b is not a unitary phenomenon but rather is a composite of activity arising from multiple intracranial sources of bioelectric activity.

Adult

Visual evoked electrical and magnetic response to half-field stimulation using pattern reversal stimulation.

The visual evoked magnetic response to half-field stimulation using pattern reversal was studied using a d.c. SQUID coupled to a second order gradiometer. The main component of the magnetic response consisted of a positive wave at around 100 ms (P100M). At the time this component was present the response to half-field stimulation consisted of an outgoing magnetic field contralateral and extending to the midline. When the left half field was stimulated the outgoing field was over the posterior right visual cortex and when the right half field was stimulated it was over the left anterior visual cortex. These findings would correctly identify a source located in the contralateral visual cortex. The orientation of the dipoles was not that previously assumed to explain the paradoxical lateralization of the visual evoked potential. The results are discussed in terms of both electrical and magnetic models of the calcarine fissure.

Adult

Colour, contrast and the visual evoked potential.

Visual evoked potentials exhibit interesting morphological changes when they are elicited by checkerboards of different spatial and chromatic contrast, counterphasing in the foveal and lower macula field. The characteristic, positive wave of the phase-reversal visual evoked potential, for example, is preceded by an increasingly prominent negative peak as luminance contrast progressively increases above 10% and, at isoluminance, the response to red and green checkerboards becomes a predominantly monophasic negative wave. To study the nature of the morphological change we synthesized these waveforms with a computer simulation consisting of Gaussian components. The amplitudes of positive and negative components were altered until the synthesized response was closely similar to the recorded data. These Gaussian components have response characteristics which are identified with those of magnocellular and parvocellular neurones.

Color Perception

VEP acuity, FPL acuity, and visual behavior of visually impaired children.

Forced-choice preferential-looking (FPL) and pattern-visual-evoked potential (VEP) acuity tests were conducted with 42 children with bilateral moderate to severe visual impairment (age, 4 months to 9 years). Within this group of children, FPL acuity testing was more successful than VEP acuity testing (98% versus 64%). Mean FPL acuity was better than mean VEP acuity (20/155 vs 20/290, P less than .002). For the children who completed both FPL and VEP acuity tests, an acuity difference score (FPL minus VEP) was derived, and compared to difference scores based on normative data. Significant differences were found between visually impaired (VI) and control difference scores (P less than .001). Difference scores of VI were larger than and opposite in sign to the control scores. Rating scores of visual behaviors supported the concurrent validity of the FPL acuities (P = .01), but not the VEP acuities.

Child

Stimulus complexity, EEG abundance gradients, and detection efficiency in a visual recognition task.

Occipital EEG was monitored while subjects inspected 27 projected patterns. The number (N) and variety (V) of elements in the patterns were varied systematically. There were three levels of N (6, 12 or 24 elements) and three levels of V (circles, squares or hexagons occupying all, one half or one third of the element locations for all levels of N). Subjects were instructed that they would be required in a post-test to recognize the patterns, among patterns which had not appeared; they were also informed that the patterns had been constructed according to a set of simple rules, but the nature of these rules was not made fully explicit. The EEG was quantified by means of low-frequency analysis, yielding measures of abundance (theta, alpha and beta) and mean dominant frequency. For the recognition task, nine stimulus items were embedded among 45 items. Recognition efficiency was measured by means of the signal detection theory discrimination index (d'). The results were as follows: (i) Both N and V were inversely related to alpha abundance (P less than 0-01); (ii) the strongest relationship between stimulus parameters and the EEG held for N and EEG beta activity (13-5-19-5 Hz;P less than 0-001), where again the EEG and N were inversely related; (iii) there was a significant (P less than 0-05) direct relationship between N and theta activity; (iv) contrary to prediction, mean dominant alpha frequency decreased as N increased; (v) d' correlated significantly with a number of effects for N, i.e. subjects who exhibited greatest EEG discriminability of items during exposure of the patterns, subsequently obtained the higher detection scores in the recognition task. The work described therefore demonstrates that only only do stimulus parameters have systematic effects upon brain activity as measured by the EEG, but that such effects have functional value and reflect aspects of efficiency. The results are fully compatible with arousal theory constructs relating physiological reactivity and performance.

Adolescent

Recognition of changes in the dimensions and categories of visual objects.

The aim of the present work is a comparison between recognition with changes in the dimensions of the objects and recognition with changes in the category of the objects. After preliminary training under tachistoscopic ocnditions, to a control set of different contour drawings are added: (1) the same drawings increased or desreased several times; (2) different new drawings with the same size. The percentage of recognized drawings is determined for such exposure time which is needed for the recognition of 60--80 per cent of the drawings in the control set. Recognition is found to be deteriorated in the case of changes both in the dimensions and in the category of the objects, the deterioration being double for the objects with changed category. This fact serves as a basis for rejecting the hypothesis that considerable changes in the dimensions could create new objects for the visual system. It was also found that recognition of pre-trained drawings is either not influenced or comparatively least influenced by the changes in the dimensions and categories. This result is probably due to the specificity of recognition of long trained objects. The results obtained do not contradict the assumption of the participation of spatial consecutive scanning of the iconic memory in the recognition of the spatial properties of visual objects.

Form Perception