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.

The properties of the motion-detecting mechanisms mediating perceived direction in stochastic displays.

Previous studies [e.g. Baker & Hess, 1998. Vision Research, 38, 1211-1222] have shown that perceived direction in displays composed of multiple, limited-lifetime, Gabor micropatterns (G) is influenced by movement both at the fine spatial scale of the internal luminance modulation (first-order motion) and the coarse spatial scale of the Gaussian, contrast window (second-order motion). However it is presently indeterminate as to whether this pattern of results is indicative of the processes by which first-order and second-order motion signals interact within the visual system per se or those by which motion information, irrespective of how it is defined, is utilised across different spatial scales. To address this issue, and more generally the properties of the mechanisms that analyse motion in such displays, we employed stochastic motion sequences composed of either G, G added to a static carrier (G + C) or G multiplied with a carrier (G*C). Crucially G*C, unlike both G and G + C, micropatterns contain no net first-order motion and second-order motion only at the scale of the internal contrast modulation. For small displacements perceived direction in all cases showed a dependence on the internal sinusoidal spatial structure of the micropatterns and characteristic oscillations were typically observed, consistent with models in which first-order motion and second-order motion are encoded on the basis of similar low-level mechanisms. Importantly for larger displacements, and also when the internal spatial structure was randomised on successive exposures (so that motion at this spatial scale was unreliable), performance tended to be veridical for all types of micropattern, even though under these conditions displacements of the G*C micropatterns should have been invisible to current, low-level, motion-detecting schemes. This suggests that both low-level motion sensors and mechanisms utilising a different motion-detecting strategy such as high-level, attentive, feature-tracking may mediate perceptual judgements in stochastic displays.

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.

Dmax for stereopsis depends on size, not spatial frequency content.

Stereoacuity depends not only on the carrier frequency of Gabor stimuli, but also upon their size. To determine if this is also the case at large disparities, we have measured the upper limit for stereopsis, "Dmax", and assessed its dependence on carrier frequency and overall envelope size. The results differ markedly from the stereoacuity data. Dmax for stereopsis is primarily dependent on the size of the envelope of the Gabor patch, and is relatively independent of its carrier frequency. These results support the proposition that stereopsis is achieved at large disparities by way of non-linear processing (envelope extraction).

Linear and non-linear filtering in stereopsis.

To better understand the spatial filtering operations underlying stereopsis, and their relationship to those underlying monocular localization of the same stimuli, we examined the dependence of stereoacuity on carrier and envelope size of Gabor patches. For stimuli of broad spatial bandwidth, stereoacuity depends on the carrier spatial frequency whereas for stimuli of narrow bandwidth, stereoacuity depends on the modulation frequency. The dependence of stereoacuity on the separation of the reference elements differs for stimuli of broad and narrow spatial frequency bandwidths. These relationships suggests that stereopsis has access to two different types of information from the early filters which we term, linear and non-linear. This distinction is important not only for understanding the relationship between monocular and stereoscopic localization, but also for understanding the different filter operations underlying stereopsis.

Binocular integration of contrast information in amblyopia.

We asked whether suppression in amblyopia could be accounted for by dichoptic masking as described in normals, operating in the presence of a contrast threshold difference between the two eyes. A dichoptic masking paradigm was employed to investigate binocular interaction in a mixed group of amblyopic subjects. Normal dichoptic masking was not seen after threshold differences between the two eyes were accounted for in the majority of subjects studied. We found that the binocular dysfunction did not merely follow as a consequence of the known monocular loss and that it depended upon the aetiology of the amblyopia and the spatial frequency of the stimulus.

Spatial information and uncertainty in anisometropic amblyopia.

Anisometropic amblyopes were found to have a reduced sensitivity for shape discrimination. The introduction of positional jitter in the elements of the display had a profound effect on the performance of the normal eye, but not on that of the amblyopic eye. On the other hand the introduction of gaussian blur affects the performance of both eyes to the same degree. We conclude that raised spatial uncertainty due to metrical scrambling is a suitable model for anisometropic amblyopia.

Is the increased spatial uncertainty in the normal periphery due to spatial undersampling or uncalibrated disarray?

There are two possible causes for the elevated positional uncertainty of the peripheral field; undersampling by post receptoral arrays or uncalibrated disarray within post receptoral arrays. In order to assess the relative influences of these two factors to peripheral spatial uncertainty, we apply an approach which has enjoyed much success in the field of colour vision to examine the trade-off between position and contrast errors for spatial vision. We show that, if our positional uncertainty in the periphery is due to spatial undersampling, a correlated contrast inaccuracy should be present. We find no evidence for this expected linkage in the peripheral field at any spatial scale. We conclude that uncalibrated neural disarray rather than undersampling is the cause of peripheral positional uncertainty.

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.