Interocular interaction of contrast and luminance signals in human primary visual cortex.

Interocular interaction in the visual system occurs under dichoptic conditions when contrast and luminance are imbalanced between the eyes. Human psychophysical investigations suggest that interocular interaction can be explained by a contrast normalization model. However, the neural processes that underlie such interactions are still unresolved. We set out to assess, for the first time, the proposed normalization model of interocular contrast interactions using magnetoencephalography (MEG) and to extend this model to incorporate interactions based on interocular luminance differences. We used MEG to record steady-state visual evoked responses (SSVER), and functional magnetic resonance imaging (fMRI) to obtain individual retinotopic maps that we used in combination with MEG source imaging in healthy participants. Binary noise stimuli were presented in monocular or dichoptic viewing and were frequency-tagged at 4 and 6 Hz. The contrast of the stimuli was modulated in a range between 0 and 32%. Monocularly, we reduced the luminance by placing a 1.5 ND filter over one eye in the maximal contrast condition. This ND filter reduces the mean light level by a factor of 30 without any alteration to the physical contrast. We observed in visual area V1 a monotonic increase in the magnitude of SSVERs with changes in contrast from 0 to 32%. For both eyes, dichoptic masking induced a decrease in SSVER signal power. This power decrease was well explained by the normalization model. Reducing mean luminance delayed monocular processing by approximately 38 ms in V1. The reduced luminance also decreased the masking ability of the eye under the filter. Predictions based on a temporal filtering model for the interocular luminance difference prior to the model's binocular combination stage were incorporated to update the normalization model. Our results demonstrate that the signals resulting from different contrast or luminance stimulation of the two eyes are combined in a way that can be explained by an interocular normalization model.

Abnormal cortical processing of pattern motion in amblyopia: evidence from fMRI.

Converging evidence from human psychophysics and animal neurophysiology indicates that amblyopia is associated with abnormal function of area MT, a motion sensitive region of the extrastriate visual cortex. In this context, the recent finding that amblyopic eyes mediate normal perception of dynamic plaid stimuli was surprising, as neural processing and perception of plaids has been closely linked to MT function. One intriguing potential explanation for this discrepancy is that the amblyopic eye recruits alternative visual brain areas to support plaid perception. This is the hypothesis that we tested. We used functional magnetic resonance imaging (fMRI) to measure the response of the amblyopic visual cortex and thalamus to incoherent and coherent motion of plaid stimuli that were perceived normally by the amblyopic eye. We found a different pattern of responses within the visual cortex when plaids were viewed by amblyopic as opposed to non-amblyopic eyes. The non-amblyopic eyes of amblyopes and control eyes differentially activated the hMT+ complex when viewing incoherent vs. coherent plaid motion, consistent with the notion that this region is centrally involved in plaid perception. However, for amblyopic eye viewing, hMT+ activation did not vary reliably with motion type. In a sub-set of our participants with amblyopia we were able to localize MT and MST within the larger hMT+ complex and found a lack of plaid motion selectivity in both sub-regions. The response of the pulvinar and ventral V3 to plaid stimuli also differed under amblyopic vs. non-amblyopic eye viewing conditions, however the response of these areas did vary according to motion type. These results indicate that while the perception of the plaid stimuli was constant for both amblyopic and non-amblyopic viewing, the network of neural areas that supported this perception was different.

Pattern-motion selective responses in MT, MST and the pulvinar of humans.

Plaid stimuli are often used to investigate the mechanisms involved in the integration and segregation of motion information. Considering the perceptual importance of such mechanisms, only a very limited number of visual brain areas have been found to be specifically involved in motion integration. These are the human (h)MT+ complex, area V3 and the pulvinar. The hMT+ complex can be functionally subdivided into two separate areas, middle temporal area (MT) and medial superior temporal area (MST); however, it is currently unclear whether these distinct sub-regions have different responses to plaid stimuli. To address this issue we used functional magnetic resonance imaging to quantify the relative response of MT and MST to component and pattern motion. Participants viewed plaid stimuli that were constrained to result in the perception of either component motion (segregation of motion information) or pattern motion (integration of motion information). MT/MST segregation was achieved using a moving dot stimulus that allowed stimulation of each visual hemifield either in unison or separately. We found pattern motion selective responses in both MT and MST. Consistent with previous reports, activity indicative of pattern motion selectivity was also found in the pulvinar as well as in other extrastriate areas. These results demonstrate that MT, MST and the pulvinar are involved in the complex motion integration mechanisms that are triggered by plaid stimuli. This reinforces the concept that integrative computations take place in a distributed neuronal circuit both in cortical and sub-cortical networks.

Binocular influences on global motion processing in the human visual system.

This study investigates four key issues concerning the binocular properties of the mechanisms that encode global motion in human vision: (1) the extent of any binocular advantage; (2) the possible site of this binocular summation; (3) whether or not purely monocular inputs exist for global motion perception; (4) the extent of any dichoptic interaction. Global motion coherence thresholds were measured using random-dot-kinematograms as a function of the dot modulation depth (contrast) for translational, radial and circular flow fields. We found a marked binocular advantage of approximately 1.7, comparable for all three types of motion and the performance benefit was due to a contrast rather than a global motion enhancement. In addition, we found no evidence for any purely monocular influences on global motion detection. The results suggest that the site of binocular combination for global motion perception occurs prior to the extra-striate cortex where motion integration occurs. All cells involved are binocular and exhibit dichoptic interactions, suggesting the existence of a neural mechanism that involves more than just simple summation of the two monocular inputs.

Only two phase mechanisms, +/-cosine, in human vision.

We evaluated the proposal that there exist detectors of the following four cardinal phases in human vision: +cosine, -cosine, +sine, and -sine. First, we assessed whether there was evidence that these cardinal phases were processed by independent 'labeled lines,' using a discrimination at detection threshold paradigm. Second, we assessed whether suprathreshold phase discrimination was best at phases intermediate between these cardinal values. Third, we tried to replicate previous evidence showing that an absence of facilitation occurs only between cosine pedestals and sine tests (or vice-versa). In all three experimental approaches we found no compelling evidence for four cardinal phase groupings. We did however find evidence for independent detectors for pure increments and decrements (+/-cosine). We suggest that phase discrimination, whether at threshold or suprathreshold, is mediated by mechanisms that encode the relative positions and contrasts of local increments and decrements within the stimulus.

The extent of the dorsal extra-striate deficit in amblyopia.

Previously, we have shown that humans with amblyopia exhibit deficits for global motion discrimination that cannot be simply ascribed to a reduction in visibility or contrast sensitivity. Deficits exist in the processing of global motion in the fronto-parallel plane that suggest reduced extra-striate function (i.e., MT) in amblyopia. Here, we ask whether such a deficit also exists for rotation and radial components of optic flow that are first processed at higher sites along the dorsal pathway (i.e., MSTd). We show that similar motion processing deficits occur in our amblyopic group as a whole for translation, rotation, and radial components of optic flow and that none of these can be solely accounted for by the reduced visibility of the stimuli. Furthermore, on a subject-by-subject basis there is no significant correlation between the motion deficits for radial and rotational motion and those for translation, consistent with independent deficits in dorsal pathway function up to and including MSTd.

Vision through an abnormal cornea: a pilot study of the relationship between visual loss from corneal distortion, corneal edema, keratoconus, and some allied corneal pathology.

Visual function was assessed by classic acuity measures as well as contrast thresholds over a wide spatial frequency range for subjects with experimentally induced corneal distortion and induced corneal edema. These results were compared with similar results for patients with uniocular keratoconus, bilateral keratoconus, and allied corneal pathology. Distortion and edema were found to produce characteristically different types of contrast attenuation at threshold (amplitude) and have quite different suprathreshold abnormalities (phase) for objects within the resolution limit and therefore could form the basis of a useful functional classification of the visual degradation from corneal pathology.

The effect of corneal edema on visual function.

Corneal edema was produced by atmospheric-induced anoxia and the visual effect was assesed by conventional acuity tests and compared with contrast sensitivity measurements. For the degree of edema which this technique produced, contrast sensitivity was depressed for only high spatial frequencies and could be considered in terms of equivalent defocus. This was not the case ofr a greater degree of edema which was simulated by a diffuser with a particle size that mimics the diffraction effects from an edematous cornea. Equivalent blur was not applicable, since low spatial frequencies were also affected and the letter and contrast sensitivity tests gave radically different results. In the light of these findings, the validity of the present letter acuity evaluation of visual function for patients in which intraocular scattering is involved is questioned.

The pattern evoked electroretinogram: its variability in normals and its relationship to amblyopia.

Electroretinograms evoked by pattern stimuli (contrast reversing gratings) were measured under steady state conditions in the normal and amblyopic eyes of 14 amblyopic individuals having Snellen acuities in the range 20/100 to 20/600. These ERGs were measured as a function of spatial frequency, and compared with the psychophysical threshold losses to the same stimuli. In all cases the authors compared the normal and fellow amblyopic eye's response while taking into account the variability of right-left eye comparisons of normal individuals for these psychophysical and electrophysiological tests. When factors such as optical focus, fixation alignment, and fixation stability have been individually optimized, no pattern ERG deficit was observed in a spatial frequency range where there were obvious psychophysical deficits to the same stimuli. Our results do not substantiate previous claims of a pattern ERG anomaly in many severely amblyopic eyes.

Contour integration and cortical processing.

Our understanding of visual processing in general, and contour integration in particular, has undergone great change over the last 10 years. There is now an accumulation of psychophysical and neurophysiological evidence that the outputs of cells with conjoint orientation preference and spatial position are integrated in the process of explication of rudimentary contours. Recent neuroanatomical and neurophysiological results suggest that this process takes place at the cortical level V1. The code for contour integration may be a temporal one in that it may only manifest itself in the latter part of the spike train as a result of feedback and lateral interactions. Here we review some of the properties of contour integration from a psychophysical perspective and we speculate on their underlying neurophysiological substrate.

The site and nature of suppression in squint amblyopia.

A significant percentage of humans have a misaligned eye (squint) due to a disruption of early visual development. In later adult life these people do not experience double vision because the visual information from their misaligned eye is actively suppressed within their visual system. Here I utilize a phenomenon called spatial adaptation which is known to have its site in the striate cortex to answer the question "is the site of suppression before, at, or after the site of adaptation?" Strabismic amblyopes who display spatial adaptation when viewing monocularly with their amblyopic eye fail to display adaptation through their amblyopic eyes under binocular viewing conditions. The lack of adaptation depends on the orientational difference between the adapting stimuli seen by each eye under binocular viewing conditions. These results suggest that suppression occurs at rather than before or after the first site of adaptation.

On the relationship between strabismic amblyopia and eccentric fixation.

Landolt C and grating acuity are compared with that normally expected of the eccentric fixation region for 10 strabismic amblyopes. The findings suggest that 2 populations of amblyopes exist. For some amblyopes visual function is that predicted of the eccentric region used for fixation, whereas for other amblyopes there is a further pathological reduction in visual function. These findings may have an important bearing on the type of orthoptic treatment used for amblyopia.

Neural recruitment explains "Weber's law" of spatial position.

We ask whether the well known Weber's law between spatial localization and element separation for high contrast, spectrally broad-band stimuli is a consequence of the organization of the early visual filters, or a fundamental constraint on the computation of spatial position by more central mechanisms. We address this question by identifying the individual contributions of mechanisms tuned to different ranges of spatial frequencies and contrast. We measure spatial-alignment and bisection error as a function of element separation at each of a number of spatial scales, using spectrally narrow-band stimuli of fixed supra-threshold contrast. We show that stimuli which minimize the extent of neural recruitment across different spatial channels before the site of extraction of the local contrast energy (and to a lesser extent across different contrast channels) do not exhibit Weber's law for either alignment or bisection. We present evidence that Weber's law for localization with increasing separation, found for stimuli of high contrast and broad-band spatial frequency content, is a consequence of the successive disengagement of unitary neural mechanisms, each of which has different spatial and contrast properties, and none of which individually exhibits Weber's law for spatial position.

Motion sensitivity and spatial undersampling in amblyopia.

The nature of the visual deficit in human amblyopia has been keenly sought over the last decade. Some confusion has arisen as to whether the motion-sensitive mechanisms known to exist in normal vision are selectively affected in humans with amblyopia. To address this issue we compare contrast thresholds for detection and direction discrimination of drifting sine-wave gratings in a group of humans with amblyopia. The results suggest that over the vast majority of the spatio-temporal range, direction of motion can be discriminated at detection threshold. Over a narrow part of the visible range there is a differential loss of sensitivity for direction discrimination over that of simple detection. However such an effect also occurs for normal vision under spatially scaled conditions and it seems likely that it is mediated by non-motion sensitive mechanisms. We show that one possible cause of this loss of direction discrimination, namely spatial undersampling within the central region of the amblyopic visual field, is not a viable explanation.

Assessment of retinal function in severely amblyopic individuals.

A quantitative psychophysical and electroretinographic assessment of severely amblyopic human eyes revealed no electrophysiological retinal analogue of the psychophysical deficit, when objectively refracted. Electroretinograms to pattern and uniform field flicker were normal in all but one amblyope. In this amblyope, although the electroretinographic deficit was uncorrelated with the psychophysical deficit, the reduced amplitude of the pattern-evoked response was also seen in the second harmonic component to uniform field stimulation. The deficit for each stimulus was only present at high signal levels. The similarity of the pattern evoked response and the second harmonic response to uniform field stimulation in normal and amblyopic eyes suggests a similar underlying process.

The contrast sensitivity gradient across the human visual field: with emphasis on the low spatial frequency range.

The regional variation of contrast sensitivity along the greater extent of each of the four principal hemi-meridia of the normal human eye was determined under photopic conditions using horizontally-orientated sinusoidal grating stimuli. The stimuli were well localized in space and frequency, and special attention was paid to the low spatial frequency range. The results confirm that contrast sensitivity is maximal for central vision for all test spatial stimuli. Extra-foveal fall-off in sensitivity can be represented as a linear function of eccentricity if the latter is expressed in relative units (i.e. periods of the stimulus). The regional variation parameter depends upon whether the horizontal or vertical field is tested and upon the spatial frequency of stimulation. The visible spatial frequency range (0.05-24 c/deg) can be approximately described by just three different rules. The fact that more than one rule is found bears upon current models of the functional organization of the visual system.

What limits the contribution of second-order motion to the perception of surface shape?

Both motion and stereopsis can be derived from contrast as well as luminance defined stimuli. It is currently assumed that these two different sources of information about objects feed into one common stage. Thus it would not be expected that their role in visual perception would be different. Here we show that although motion can be carried by contrast-defined elements, such motion is not used to define three-dimensional (3D) surfaces. A similar effect has been reported in stereopsis; although such contrast-defined elements can give signed disparity signals they nevertheless do not contribute to the percept of shape. We show that the reason for this lies in the inability of the second order signals to cohere or bind across space/spatial scales rather than a characteristic of the elementary motion signals per se.

The spatial localization deficit in amblyopia.

There have now been numerous reports of a spatial localization deficit in amblyopia but none so far have tackled (1) the relationship between the contrast sensitivity and spatial localization deficits and (2) whether the spatial localization deficit is best described in units of visual angle or in terms of the underlying filter size. These issues are germane because they lie at the very heart of our understanding of the underlying deficit in amblyopia. To answer these questions we use spatially bandpass stimuli so that we can readily compare detection and localization for the same stimuli at each of a number of spatial scales. For some amblyopes (all strabismics and a minority of anisometropes) the contrast sensitivity defect neither underlies nor covaries with the spatial localization deficit. In the majority of anisometropic amblyopes, the contrast sensitivity loss is a complete description. The spatial localization deficit in amblyopia is of two independent kinds; positional inaccuracy and positional distortion. The positional inaccuracy deficit which can occur in varying degrees in both strabismic and anisometropic amblyopia, affects all spatial scales equally and therefore is best thought of in terms of a constant fraction of the underlying filter size in the space-frequency plane. The positional distortion deficit which can also occur to varying degrees in both strabismic and anisometropic forms can not be easily understood within this metric at least for strabismics.

Temporal properties of human visual filters: number, shapes and spatial covariation.

The temporal properties of the foveal visual filters were revealed using a method which is a variant on previously used noise masking paradigms. This enables the temporal properties of the mechanisms underlying threshold detection of a spatio-temporal probe to be measured. In accord with recent suggestions these results support the existence of three temporal mechanisms. The evidence for the third, higher temporal mechanism is only persuasive at low spatial frequencies. Furthermore, the results suggest that although there is some degree of spatio-temporal covariation in the filtering properties either of individual filters or across the filter population, the well known spatio-temporal covariation in human detection sensitivity is adequately explained by a sensitivity scaling of individual temporal filters with approximately invariant temporal properties.

Temporal frequency filters in the human peripheral visual field.

The temporal filtering properties of the human peripheral field were investigated by means of measuring: (1) modulation transfer functions for a range of spatial frequencies at four visual field locations (0, 10, 30 and 50 degrees), (2) the contrast of a masking stimulus required to extinguish the visibility of just suprathreshold probes. Results suggest that the number of temporal filters governing detection threshold is dependent upon both eccentricity and spatial frequency. For near-foveal viewing three temporal filters were found (one low-pass and two band-pass), whereas at far eccentricities only one was found (band-pass). A similar result was obtained by modeling the modulation transfer function by simply scaling the sensitivities of three independently derived filters. Our data suggest that (1) changes in the modulation transfer function with respect to spatial frequency and eccentricity can be adequately explained by the changes in sensitivity of a small number of spatio-temporal separable filters; (2) the peripheral field is not merely a coarser version of the fovea but has qualitative differences which may be thought to emphasize the transient properties of the stimulus.