Induced gamma activity in primary visual cortex is related to luminance and not color contrast: an MEG study

Gamma activity in the visual cortex has been reported in numerous EEG studies of coherent and illusory figures. A dominant theme of many such findings has been that temporal synchronization in the gamma band in response to these identifiable percepts is related to perceptual binding of the common features of the stimulus. In two recent studies using magnetoencephalography (MEG) and the beamformer analysis technique, we have shown that the magnitude of induced gamma activity in visual cortex is dependent upon independent stimulus features such as spatial frequency and contrast. In particular, we showed that induced gamma activity is maximal in response to gratings of 3 cycles per degree (3 cpd) of high luminance contrast. In this work, we set out to examine stimulus contrast further by using isoluminant red/green gratings that possess color but not luminance contrast using the same cohort of subjects. We found no induced gamma activity in V1 or visual cortex in response to the isoluminant gratings in these subjects who had previously shown strong induced gamma activity in V1 for luminance contrast gratings.

Fundamental characteristics of space-charge waves in photorefractive sillenite crystals

We investigate experimentally the fundamental characteristics of space-charge waves excited in a photorefractive crystal of Bi12SiO20. Features such as their transient rise and decay as well as their steady-state frequency response are investigated. Based on this, we find the dependence of the space-charge waves' quality factor on spatial frequency and electric-field biasing. The experimental findings are compared with the linear space-charge wave theory developed previously by Sturman et al. [J. Opt. Sec. Am. B 10, 1919 (1993)].

Contextual modulation involves suppression and facilitation from the center and the surround

In psychophysics, cross-orientation suppression (XOS) and cross-orientation facilitation (XOF) have been measured by investigating mask configuration on the detection threshold of a centrally placed patch of sine-wave grating. Much of the evidence for XOS and XOF comes from studies using low and high spatial frequencies, respectively, where the interactions are thought to arise from within (XOS) and outside (XOF) the footprint of the classical receptive field. We address the relation between these processes here by measuring the effects of various sizes of superimposed and annular cross-oriented masks on detection thresholds at two spatial scales (1 and 7 c/deg) and on contrast increment thresholds at 7 c/deg. A functional model of our results indicates the following (1) XOS and XOF both occur for superimposed and annular masks. (2) XOS declines with spatial frequency but XOF does not. (3) The spatial extent of the interactions does not scale with spatial frequency, meaning that surround-effects are seen primarily at high spatial frequencies. (4) There are two distinct processes involved in XOS: direct divisive suppression and modulation of self-suppression. (5) Whether XOS or XOF wins out depends upon their relative weights and mask contrast. These results prompt enquiry into the effect of spatial frequency at the single-cell level and place new constraints on image-processing models of early visual processing. © ARVO.

Remote facilitation in the Fourier domain

To explore spatial interactions between visual mechanisms in the Fourier domain we measured detection thresholds for vertical and horizontal sine-wave gratings (4.4 deg diameter) over a range of spatial frequencies (0.5-23 c/deg) in the presence of grating and plaid masks with component contrasts of 8%, orientations of ±45° and a spatial frequency of 1 c/deg. The mask suppressed the target grating over a range of ±1 octave, and the plaid produced more suppression than the grating, consistent with summation of mask components in a broadly tuned contrast gain pool. At greater differences in spatial frequency (∼3 octaves), the plaid and grating masks both produced about 3 dB of facilitation (they reduced detection thresholds by a factor of about √2). At yet further distances (∼4 octaves) the masks had no effect. The facilitation cannot be attributed to a reduction of uncertainty by the mask because (a) it occurs for mask components that have very different spatial frequencies and orientations from the test and (b) the large stimulus size and central fixation point mean there was no spatial uncertainty that could be reduced. We suggest the results are due to long-range sensory interactions (in the Fourier domain) between mask and test-channels. The effects could be due to either direct facilitation or disinhibition. © 2006 Elsevier Ltd. All rights reserved.

Interocular suppression is gated by interocular feature matching

We present a new form of contrast masking in which the target is a patch of low spatial frequency grating (0.46 c/deg) and the mask is a dark thin ring that surrounds the centre of the target patch. In matching and detection experiments we found little or no effect for binocular presentation of mask and test stimuli. But when mask and test were presented briefly (33 or 200 ms) to different eyes (dichoptic presentation), masking was substantial. In a 'half-binocular' condition the test stimulus was presented to one eye, but the mask stimulus was presented to both eyes with zero-disparity. This produced masking effects intermediate to those found in dichoptic and full-binocular conditions. We suggest that interocular feature matching can attenuate the potency of interocular suppression, but unlike in previous work (McKee, S. P., Bravo, M. J., Taylor, D. G., & Legge, G. E. (1994) Stereo matching precedes dichoptic masking. Vision Research, 34, 1047) we do not invoke a special role for depth perception. © 2004 Elsevier Ltd. All rights reserved.

Grating and plaid masks indicate linear summation in a contrast gain pool

In human vision, the response to luminance contrast at each small region in the image is controlled by a more global process where suppressive signals are pooled over spatial frequency and orientation bands. But what rules govern summation among stimulus components within the suppressive pool? We addressed this question by extending a pedestal plus pattern mask paradigm to use a stimulus with up to three mask components: a vertical 1 c/deg pedestal, plus pattern masks made from either a grating (orientation = -45°) or a plaid (orientation = ±45°), with component spatial frequency of 3 c/deg. The overall contrast of both types of pattern mask was fixed at 20% (i.e., plaid component contrasts were 10%). We found that both of these masks transformed conventional dipper functions (threshold vs. pedestal contrast with no pattern mask) in exactly the same way: The dipper region was raised and shifted to the right, but the dipper handles superimposed. This equivalence of the two pattern masks indicates that contrast summation between the plaid components was perfectly linear prior to the masking stage. Furthermore, the pattern masks did not drive the detecting mechanism above its detection threshold because they did not abolish facilitation by the pedestal (Foley, 1994). Therefore, the pattern masking could not be attributed to within-channel masking, suggesting that linear summation of contrast signals takes place within a suppressive contrast gain pool. We present a quantitative model of the effects and discuss the implications for neurophysiological models of the process. © 2004 ARVO.

Area summation and masking

At detection threshold, sensitivity improves as the area of a test grating increases, but not when the test is placed on a pedestal and the task becomes contrast discrimination (G. E. Legge, & J. M. Foley, 1980). This study asks whether the abolition of area summation is specific to the situation where mask and test stimuli have the same spatial frequency and orientation ("within-channel" masking) or is more general, also occurring when mask and test stimuli are very different ("cross-channel" masking). Threshold versus contrast masking functions were measured where the test and mask were either both small (SS), both large (LL), or small and large, respectively (SL). For within-channel masking, facilitation and area summation were found at low mask contrasts, but the results for SS and LL converged at intermediate contrasts and above, replicating Legge and Foley (1980). For all three observers, less facilitation was found for SL than for SS. For cross-channel masking, area summation occurred across the entire masking function and results for SS and SL were identical. The results for the entire data set were well fit by an extended version of a contrast masking model (J. M. Foley, 1994) in which the weights of excitatory and suppressive surround terms were free parameters. I conclude that (i) there is no empirical abolition of area summation for cross-channel masking, (ii) within-channel area summation can be abolished empirically without being disabled in the model, (iii) observers are able to restrict the area of spatial integration, but not suppression, (iv) extending a cross-channel mask to the surround has no effect on contrast detection, and (v) there is a formal similarity between area summation and contrast adaptation. © 2004 ARVO.

Extrinsic factors in the perception of bistable motion stimuli

When viewing a drifting plaid stimulus, perceived motion alternates over time between coherent pattern motion and a transparent impression of the two component gratings. It is known that changing the intrinsic attributes of such patterns (e.g. speed, orientation and spatial frequency of components) can influence percept predominance. Here, we investigate the contribution of extrinsic factors to perception; specifically contextual motion and eye movements. In the first experiment, the percept most similar to the speed and direction of surround motion increased in dominance, implying a tuned integration process. This shift primarily involved an increase in dominance durations of the consistent percept. The second experiment measured eye movements under similar conditions. Saccades were not associated with perceptual transitions, though blink rate increased around the time of a switch. This indicates that saccades do not cause switches, yet saccades in a congruent direction might help to prolong a percept because i) more saccades were directionally congruent with the currently reported percept than expected by chance, and ii) when observers were asked to make deliberate eye movements along one motion axis, this increased percept reports in that direction. Overall, we find evidence that perception of bistable motion can be modulated by information from spatially adjacent regions, and changes to the retinal image caused by blinks and saccades.

The influence of age on the pattern and flash visual evoked field

We have investigated the effect of ageing on the visual system using the relatively new technique of magentoencephalography (MEG). This technique measures the magnetic signals produced by the visual system using a SQUID magnetometer. The magnetic visual evoked field (VEF) was measured over the occipital cortex to pattern and flash stimuli in 86 normal subjects aged 15 - 86 years. Factors that influenced subject defocussing or defixating the stimulus or selective attention were controlled as far as possible. The latency of the major positive component to the pattern reversal stimulus (P100M) increased with age particularly after the age of 55 years while the amplitude of the P100M decreased over the life span. The latency of the major flash component (P2M) increased much more slowly with age, while its amplitude decreased in only a proportion of elderly subjects. Changes in the P100M with age may reflect senile changes in the eye and optic nerve, e.g. senile miosis or degenerative changes in the retina. The P2M may be more susceptible to senile changes in the retina. The data suggest that the spatial frequency channels deteriorate more rapidly with age than the luminance channels and that MEG may be an effective method of studying ageing in the visual system.

Fundamental characteristics of space-charge waves in photorefractive sillenite crystals

We investigate experimentally the fundamental characteristics of space-charge waves excited in a photorefractive crystal of Bi12SiO20. Features such as their transient rise and decay as well as their steady-state frequency response are investigated. Based on this, we find the dependence of the space-charge waves' quality factor on spatial frequency and electric-field biasing. The experimental findings are compared with the linear space-charge wave theory developed previously by Sturman et al. [J. Opt. Sec. Am. B 10, 1919 (1993)].

Further studies of the visually evoked subcortical potential in man

The Visually Evoked Subcortical Potential, a far-field signal, was originally defined to flash stimulation as a triphasic positive-negative-positive complex with mean latencies of P21 N26.2 P33.6 (Harding and Rubinstein 1980). Inconsistent with its subcortical source however, the signal was found to be tightly localised to the mastoid. This thesis re-examines the earlier protocols using flash stimulation and with auditory masking establishes by topographic studies that the VESP has a widespread scalp distribution, consistent with a far-field source of the signal, and is not a volume-conducted electroretinogram (ERG). Furthermore, mastoid localisation indicates auditory contamination from the click, on discharge of the photostimulator. The use of flash stimulation could not precisely identify the origin of the response. Possible sources of the VESP are the lateral geniculate body (LGB) and the superior colliculus. The LGB received 80% of the nerve fibres from the retina, and responds to high contrast achromatic stimulation in the form of drifting gratings of high spatial frequencies. At low spatial frequencies, it is more sensitive to colour. The superior colliculus is insensitive to colour and suppressed by contrast and responds to transitory rapid movements, and receives about 20% of the optic nerve fibres. A pattern VESP was obtained to black and white checks as a P23.5 N29.2 P34 complex in 93% of normal subjects at an optimal check size of 12'. It was also present as a P23.0 N28.29 P32.23 complex to red and green luminance balanced checks at 2o check size in 73% of subjects. These results were not volume-conducted pattern electroretinogram responses. These findings are consistent with the spatial frequency properties of the lateral geniculate body which is the considered source of the signal. With further work, the VESP may supplement electrodiagnosis of post-chiasmal lesions.

Contrast matching and discrimation in human vision

The aim of this work was to investigate human contrast perception at various contrast levels ranging from detection threshold to suprathreshold levels by using psychophysical techniques. The work consists of two major parts. The first part deals with contrast matching, and the second part deals with contrast discrimination. Contrast matching technique was used to determine when the perceived contrasts of different stimuli were equal. The effects of spatial frequency, stimulus area, image complexity and chromatic contrast on contrast detection thresholds and matches were studied. These factors influenced detection thresholds and perceived contrast at low contrast levels. However, at suprathreshold contrast levels perceived contrast became directly proportional to the physical contrast of the stimulus and almost independent of factors affecting detection thresholds. Contrast discrimination was studied by measuring contrast increment thresholds which indicate the smallest detectable contrast difference. The effects of stimulus area, external spatial image noise and retinal illuminance were studied. The above factors affected contrast detection thresholds and increment thresholds measured at low contrast levels. At high contrast levels, contrast increment thresholds became very similar so that the effect of these factors decreased. Human contrast perception was modelled by regarding the visual system as a simple image processing system. A visual signal is first low-pass filtered by the ocular optics. This is followed by spatial high-pass filtering by the neural visual pathways, and addition of internal neural noise. Detection is mediated by a local matched filter which is a weighted replica of the stimulus whose sampling efficiency decreases with increasing stimulus area and complexity. According to the model, the signals to be compared in a contrast matching task are first transferred through the early image processing stages mentioned above. Then they are filtered by a restoring transfer function which compensates for the low-level filtering and limited spatial integration at high contrast levels. Perceived contrasts of the stimuli are equal when the restored responses to the stimuli are equal. According to the model, the signals to be discriminated in a contrast discrimination task first go through the early image processing stages, after which signal dependent noise is added to the matched filter responses. The decision made by the human brain is based on the comparison between the responses of the matched filters to the stimuli, and the accuracy of the decision is limited by pre- and post-filter noises. The model for human contrast perception could accurately describe the results of contrast matching and discrimination in various conditions.

Neuromagnetic investigations of functional organisation within human visual cortex

This thesis describes a series of experimental investigations into the functional organisation of human visual cortex using neuromagnetometry.This technique combines good spatial and temporal resolution enabling identification of the location and temporal response characteristics of cortical neurones within alert humans. To activate different neuronal populations and cortical areas a range of stimuli were used, the parameters of which were selected to match the known physiological properties of primate cortical neurones. In one series of experiments the evoked magnetic response was recorded to isoluminant red/green gratings. Co-registration of signal and magnetic resonance image data indicated a contribution to the response from visual areas V1, V2 and V4. To investigate the spatio-temporal characteristics of neurones within area V1 the evoked response was recorded for a range of stimulus spatial and temporal frequencies. The response to isoluminant red/green gratings was dominated by a major component which was found to have bandpass spatial frequency tuning with a peak at 1-2 cycles/degree, falling to the level of the noise at 6-8 cycles/degree. The temporal frequency tuning characteristics of the response showed bimodal sensitivity with peaks at 0-1Hz and 4Hz. In a further series of experiments the luminance evoked response was recorded to red/black, yellow/black and achromatic gratings and in all cases was found to be more complex than the isoluminant chromatic response, comprising up to three distinct components. The major response peak showed bandpass spatial frequency tuning characteristics, peaking at 6-8 cycles/degree, falling to the level of the noise at 12-16 cycles/degree. The results provide evidence to suggest that within area V1 the same neuronal population encodes both chromatic and luminance information and has spatial frequency tuning properties consistent with single-opponent cells. Furthermore, the results indicate that cells within area V1 encode chromatic motion information over a wide range of temporal frequencies with temporal response characteristics suggestive of the existence of a sub-population of cells sensitive to high temporal frequencies.

Investigations into visual hyperacuity

Visual hyperacuities.are a group of thresholds whose values surpass that expected by the anatomical and optical constraints of the eye. There are many variables which affect hyperacuities of which this thesis considers the following .. 1. The effect of contrast on displacement detection and bisection acuity. It is proposed that spatial summation may account for the different response of these two hyperacuities compared with the contrast response of vernier acuity. 2. The effect of references on displacement detection. These were shown to greatly enhance performance when present. Their effect was, however, dependent upon the temporal characteristics of the displacement. 3. The effect of spatial frequency on vernier acuity. Evidence from this experiment suggests that vernier performance can be explained on the basis of the output of orientationally selective spatial frequency filters. 4. Evidence for a weighting function for visual location using random dot clusters. The weighting attached to different parts of the retinal light distribution was found to alter non-linearly with increasing offset from the geometric center of the cluster. A relationship between dot density and peak amplitude of the weighting function was found. 5. Spatial scaling of vernier acuity in the peripheral field. With careful choice of a technique which did not allow separation and eccentricity to co-vary it was found possible to scale vernier acuity both for two lines and two separated dots. 6. The effect of increasing age on hyperacuity. No change in vernier acuity with age was found which contrasted with displacement detection and bisection acuity both of which showed a significant decline with increasing age.

A reevaluation of achromatic spatio-temporal vision: nonoriented filters are monocular, they adapt, and can be used for decision making at high flicker speeds

Masking, adaptation, and summation paradigms have been used to investigate the characteristics of early spatio-temporal vision. Each has been taken to provide evidence for (i) oriented and (ii) nonoriented spatial-filtering mechanisms. However, subsequent findings suggest that the evidence for nonoriented mechanisms has been misinterpreted: those experiments might have revealed the characteristics of suppression (eg, gain control), not excitation, or merely the isotropic subunits of the oriented detecting mechanisms. To shed light on this, we used all three paradigms to focus on the ‘high-speed’ corner of spatio-temporal vision (low spatial frequency, high temporal frequency), where cross-oriented achromatic effects are greatest. We used flickering Gabor patches as targets and a 2IFC procedure for monocular, binocular, and dichoptic stimulus presentations. To account for our results, we devised a simple model involving an isotropic monocular filter-stage feeding orientation-tuned binocular filters. Both filter stages are adaptable, and their outputs are available to the decision stage following nonlinear contrast transduction. However, the monocular isotropic filters (i) adapt only to high-speed stimuli—consistent with a magnocellular subcortical substrate—and (ii) benefit decision making only for high-speed stimuli (ie, isotropic monocular outputs are available only for high-speed stimuli). According to this model, the visual processes revealed by masking, adaptation, and summation are related but not identical.