Anthony M. Norcia

Collinear stimuli regulate visual responses depending on cell's contrast threshold

Neurons in the primary visual cortex are selective for the size, orientation and direction of motion of patterns falling within a restricted region of visual space known as the receptive field. The response to stimuli presented within the receptive field can be facilitated or suppressed by other stimuli falling outside the receptive field which, when presented in isolation, fail to activate the cell. Whether this interaction is facilitative,,, or suppressive,,,, depends on the relative orientation of pattern elements inside and outside the receptive field. Here we show that neuronal facilitation preferentially occurs when a near-threshold stimulus inside the receptive field is flanked by higher-contrast, collinear elements located in surrounding regions of visual space. Collinear flanks and orthogonally oriented flanks, however, both act to reduce the response to high-contrast stimuli presented within the receptive field. The observed pattern of facilitation and suppression may be the cellular basis for the observation in humans that the detectability of an oriented pattern is enhanced by collinear flanking elements. Modulation of neuronal responses by stimuli falling outside their receptive fields may thus represent an early neural mechanism for encoding objects and enhancing their perceptual saliency.

Spatial frequency sweep VEP : visual acuity during the first year of life

The grating acuity of 197 infants from 1 week to 53 weeks of age was measured using the visual evoked potential (VEP) in response to counterphase grating stimulation. The gratings were presented as a 10 sec spatial frequency sweep which spanned the acuity limit. The amplitude and phase of the second harmonic response were extracted by discrete Fourier analysis. The VEP amplitude versus spatial frequency function showed narrow spatial frequency tuning with amplitude peaks at one or more spatial frequencies. The phase of the response at medium to high spatial frequencies was generally constant at a spatial frequency peak, followed by a progressive phase lag with increasing spatial frequency. Grating acuity was estimated by linear extrapolation to zero microvolts of the highest spatial frequency peak in the VEP amplitude versus spatial frequency function. This visual acuity estimate increased from a mean of 4.5 c/deg during the first month to about 20 c/deg at 8-13 months of age. The VEP acuities at 1 month are a factor of three to five higher than previously reported for pattern reversal or pattern appearance stimuli. By 8 months VEP grating resolution was not reliably different from adult levels in the same apparatus.

Event-related brain potentials to human faces in infants.

Event-related brain potentials (ERPs) in response to tachistoscopically presented photographs of 2 human faces were recorded for 4--7-month-old infants. For each infant 1 face was chosen to be presented frequently (p = .88, a low-information event) and the other infrequently (p = .12, a high-information event). Both types of events elicited in our infants a long-latency negative ERP wave (ca. 700 msec), termed Nc, and a long-latency positive wave (ca. 1,360 msec), termed Pc. We found that the discrepant, infrequently presented face elicited Nc waves which were higher in amplitude and longer in latency than those elicited by the frequent face. These differences suggest that our infants were able to remember the frequently presented face from trial to trial and to discriminate it from the discrepant face. The discrepant event elicited Pc waves which were insignificantly higher in amplitude than those elicited by frequent events. In adults and children, discrepant events have been found by numerous researchers to elicit positive P3 waves (latency ca. 300--800 msec). In our study, however, such waves could not be discerned. So, of all of the ERP waves which have been related to cognitive processes, the wave which is maturationally the earliest to appear is the Nc wave, which has been related to the perception of attention-getting events or events of interest to the subject. Our findings suggest that ERP responses could provide a sensitive means for investigating infant cognitive development since they do not depend upon an integrated motor-response system.

Development of contrast sensitivity in the human infant

Contrast sensitivity and grating acuity were measured using the sweep VEP method in a group of 48 infants from 2 to 40 weeks of age and in a group of 10 adults. Sinusoidal gratings were reversed in contrast at 12 alternations per sec at a space-average luminance of 220 cd/m2. During 10 sec trials, either the contrast or the spatial frequency was increased in a series of 19 steps. Thresholds were estimated by extrapolation of the VEP response functions to zero amplitude. The contrast threshold at low spatial frequencies developed rapidly from 7% contrast at 2-3 weeks to an asymptote of 0.5% at 9 weeks. For adults, maximum sensitivity at low spatial frequencies was 0.32-0.22%. The sweep VEP estimate of grating acuity showed a gradual increase in spatial frequency with age, starting at 5 c/deg during the first month and reaching 16.3 c/deg at 8 months. The mean adult acuity was 31.9 c/deg. There appeared to be two phases in the development of contrast sensitivity and acuity. Between 4 and 9 weeks overall contrast sensitivity increased by a factor of 4-5 at all spatial frequencies. Beyond 9 weeks, contrast sensitivity at low spatial frequencies remained constant, while sensitivity increased systematically at higher spatial frequencies.

The steady-state visual evoked potential in vision research: A review

Periodic visual stimulation and analysis of the resulting steady-state visual evoked potentials were first introduced over 80 years ago as a means to study visual sensation and perception. From the first single-channel recording of responses to modulated light to the present use of sophisticated digital displays composed of complex visual stimuli and high-density recording arrays, steady-state methods have been applied in a broad range of scientific and applied settings.The purpose of this article is to describe the fundamental stimulation paradigms for steady-state visual evoked potentials and to illustrate these principles through research findings across a range of applications in vision science.

Neurophysiological Evidence for Contrast Dependent Long-range Facilitation and Suppression in the Human Visual Cortex

Long-range spatial interactions in human visual cortex were explored using a lateral masking paradigm. Visual evoked potentials (VEPs) elicited by a Gabor signal presented in isolation or in the presence of two flanking high-contrast Gabor signals (masks) were measured. Response amplitude and phase were recorded for a vertically oriented test, for horizontal and vertical masks and for combinations of vertical tests and vertical or horizontal masks. The amplitudes and phases of the test alone and mask alone responses were added coherently to predict the amplitude for collinear and orthogonal lateral masking conditions. Additivity failures were taken as evidence for neural interactions. At a target-to-mask distance of 2 deg, VEP amplitude exceeded the linear prediction for test contrasts in the range of 8-16% for the collinear, co-axial target/mask combination. Measured response phase also led predicted response phase over the same range of contrast. The VEP amplitudes were less than the linear prediction in the orthogonal target/mask combination and measured response phase lagged the predicted phase. Significant facilitation occurred with collinear test/mask combinations up to at least 3 deg of separation (nine wavelengths). Co-oriented, but non-collinear test/mask combinations (oblique test and mask, horizontal test and mask) did not produce facilitation. Contrast gain thus appears to be set over considerable distances in a configuration-specific fashion.

Measurement of spatial contrast sensitivity with the swept contrast VEP

Contrast response functions (CRFs) for the VEP were obtained with a Discrete Fourier Transform (DFT) technique employing swept contrast gratings. VEP CRFs in infants were found to have a form similar to those observed in adults, being linear functions of log contrast over a range of near-threshold contrasts. CRFs with low and high contrast lobes were present in infants, as they are in adults. Contrast thresholds were estimated by extrapolation of the CRF to zero microvolts. The effects of additive EEG noise and of the DFT data window on the shape of the measured CRF are considered. For large signals, the measured CRF is nearly independent of the additive noise, but at small signal values additive noise introduces a small bias towards larger amplitudes. The VEP signal-plus-noise distribution was modeled as a family of Rice distributions in order to evaluate the effects of bias on the estimates of threshold. The amount of bias depends inversely upon the slope of the CRF. The amount of bias introduced by a smoothing window also depends upon slope of the CRF as well as the sweep rate. The combined effects of additive noise and window bias were such that the total bias was nearly independent of CRF slope. Sweep VEP contrast thresholds were shown empirically to be unaffected by changes in the range of contrast swept.

Abnormal long-range spatial interactions in amblyopia.

Neural interactions between widely separated stimuli were explored with psychophysical and visual evoked potential (VEP) measures in normal and amblyopic observers. Contrast detection thresholds were measured psychophysically for small foveally viewed Gabor patches presented in isolation and in the presence of similar, but laterally displaced flanks. The amplitude and phase of VEPs elicited by similar targets were also measured. The presence of neural interaction between the target and flank responses was assessed by comparing the unflanked threshold to the flanked threshold in the psychophysical experiments and by comparing the response predicted by the algebraic sum of test and flank responses to that measured when test and flanks were presented simultaneously. In normal observers simultaneous presentation of test and flank targets produces a VEP response that is up to a factor of two larger than the linear prediction (facilitation). Psychophysical threshold is also facilitated by a comparable factor. Facilitation was found mainly for configurations in which local (carrier) and global (patch) orientations resulted in collinearity, independent of global orientation (meridian). Amblyopic observers showed several deviations from the normal pattern. The facilitation for the collinear configurations was either markedly lower than normal or was replaced by inhibition. The normal pattern of spatial interaction may facilitate the grouping of collinear line segments into smooth curves. In contrast, abnormal long-range spatial interactions may underlie the grouping disorders and perceptual distortions found in amblyopia.

A new test of contour integration deficits in patients with a history of disrupted binocular experience during visual development

Previous studies have suggested that the integration of orientation information across space is impaired in amblyopia. We developed a method for quantifying orientation-domain processing using a test format that is suitable for clinical application. The test comprises a graded series of cards where each card includes a closed path (contour) of high contrast Gabor signals embedded in a random background of Gabor signals. Contour visibility in both normals and patients with histories of abnormal binocular vision depends jointly on the spacing of elements on the contour as well as background element density. Strabismic amblyopes show significant degradation of performance compared to normals. Small but significant losses in sensitivity were also observed in a group of non-amblyopic strabismus patients. Threshold measurements made with contrast reducing diffusers indicated that the amblyopic loss is not due to the reduced contrast sensitivity of the amblyopic eye. An abnormal pattern of long-range connectivity between spatial filters or a loss of such connectivity appears to be the primary source of contour integration deficits in amblyopia and strabismus.

Cue-invariant networks for figure and background processing in human visual cortex

Lateral occipital cortical areas are involved in the perception of objects, but it is not clear how these areas interact with first tier visual areas. Using synthetic images portraying a simple texture-defined figure and an electrophysiological paradigm that allows us to monitor cortical responses to figure and background regions separately, we found distinct neuronal networks responsible for the processing of each region. The figure region of our displays was tagged with one temporal frequency (3.0 Hz) and the background region with another (3.6 Hz). Spectral analysis was used to separate the responses to the two regions during their simultaneous presentation. Distributed source reconstructions were made by using the minimum norm method, and cortical current density was measured in a set of visual areas defined on retinotopic and functional criteria with the use of functional magnetic resonance imaging. The results of the main experiments, combined with a set of control experiments, indicate that the figure region, but not the background, was routed preferentially to lateral cortex. A separate network extending from first tier through more dorsal areas responded preferentially to the background region. The figure-related responses were mostly invariant with respect to the texture types used to define the figure, did not depend on its spatial location or size, and mostly were unaffected by attentional instructions. Because of the emergent nature of a segmented figure in our displays, feedback from higher cortical areas is a likely candidate for the selection mechanism by which the figure region is routed to lateral occipital cortex.