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.

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.

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.

Restoration of binocular vision in amblyopia.

PURPOSE: To develop a treatment for amblyopia based on re-establishing binocular vision. METHODS: A novel procedure is outlined for measuring and reducing the extent to which the fixing eye suppresses the fellow amblyopic eye in adults with amblyopia. We hypothesize that suppression renders a structurally binocular system, functionally monocular. RESULTS: We demonstrate that strabismic amblyopes can combine information normally between their eyes under viewing conditions where suppression is reduced by presenting stimuli of different contrast to each eye. Furthermore we show that prolonged periods of binocular combination leads to a strengthening of binocular vision in strabismic amblyopes and eventual combination of binocular information under natural viewing conditions (stimuli of the same contrast in each eye). Concomitant improvement in monocular acuity of the amblyopic eye occurs with this reduction in suppression and strengthening of binocular fusion. Additionally, stereoscopic function was established in the majority of patients tested. We have implemented this approach on a headmounted device as well as on a handheld iPod. CONCLUSION: This provides the basis for a new treatment of amblyopia, one that is purely binocular and aimed at reducing suppression as a first step.

A new binocular approach to the treatment of amblyopia in adults well beyond the critical period of visual development.

PURPOSE: The present treatments for amblyopia are predominantly monocular aiming to improve the vision in the amblyopic eye through either patching of the fellow fixing eye or visual training of the amblyopic eye. This approach is problematic, not least of which because it rarely results in establishment of binocular function. Recently it has shown that amblyopes possess binocular cortical mechanisms for both threshold and suprathreshold stimuli. METHODS: We outline a novel procedure for measuring the extent to which the fixing eye suppresses the fellow amblyopic eye, rendering what is a structurally binocular system, functionally monocular. RESULTS: Here we show that prolonged periods of viewing (under the artificial conditions of stimuli of different contrast in each eye) during which information from the two eyes is combined leads to a strengthening of binocular vision in strabismic amblyopes and eventual combination of binocular information under natural viewing conditions (stimuli of the same contrast in each eye). Concomitant improvement in monocular acuity of the amblyopic eye occurs with this reduction in suppression and strengthening of binocular fusion. Furthermore, in a majority of patients tested, stereoscopic function is established. CONCLUSIONS: This provides the basis for a new treatment of amblyopia, one that is purely binocular and aimed at reducing suppression as a first step.

Measurement of suprathreshold binocular interactions in amblyopia.

It has been established that in amblyopia, information from the amblyopic eye (AME) is not combined with that from the fellow fixing eye (FFE) under conditions of binocular viewing. However, recent evidence suggests that mechanisms that combine information between the eyes are intact in amblyopia. The lack of binocular function is most likely due to the imbalanced inputs from the two eyes under binocular conditions [Baker, D. H., Meese, T. S., Mansouri, B., & Hess, R. F. (2007b). Binocular summation of contrast remains intact in strabismic amblyopia. Investigative Ophthalmology & Visual Science, 48(11), 5332-5338]. We have measured the extent to which the information presented to each eye needs to differ for binocular combination to occur and in doing so we quantify the influence of interocular suppression. We quantify these suppressive effects for suprathreshold processing of global stimuli for both motion and spatial tasks. The results confirm the general importance of these suppressive effects in rendering the structurally binocular visual system of a strabismic amblyope, functionally monocular.

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.

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.

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.

How many positions can we perceptually encode, one or many?

Here we show that our sensitivity for discriminating relative position across the visual field is limited. In experiment 1 we show that we are much worse at detecting a texture defined by the relative position of elements within an array than would be expected if we had access to multiple estimates of relative position across the visual field. In experiment 2 we show that human performance is impaired for positional judgments when there is uncertainty as to which of a number of possible elements is misaligned. This impairment is greater than one would expect from an ideal observer model and greater than that found for a comparable task involving orientation. It is consistent with positional thresholds being determined by only one estimate of relative position. In experiment 3 we estimate the number of suprathreshold positional signals that can be pooled at the same time across the visual field using a standard summation variance paradigm. The results suggest that the human visual system is limited to one estimate of position, but additional estimates can be built up serially over time; however, this process is slow and probably cognitive in nature. These experiments taken as a whole suggest that only one estimate of relative position (i.e. relative to a predefined reference) at a time is accessible at the perceptual level.

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.

Temporal detection in human vision: dependence on eccentricity.

Studies of human perception of time-varying luminance often aim to estimate either temporal impulse response shapes or temporal modulation transfer functions (MTFs) of putative temporal processing mechanisms. Previously, temporal masking data have been used to estimate the properties and numbers of these temporal mechanisms in central vision for 1 cycle per degree (cpd) targets [Fredericksen and Hess (1998)]. The same methods have been used to explore how these properties change with stimulus energy [Fredericksen and Hess (1997)] and spatial frequency [Fredericksen and Hess (1999)]. We present here analyses of the properties of temporal mechanisms that detect temporal variations of luminance in peripheral vision. The results indicate that a two-filter model provides the best model for our masking data, but that no multiple filter model provides an acceptable fit across the range of parameters varied in this study. Single-filter modelling shows differences between processing mechanisms at 1 cpd in central vision and those that operate eccentrically. We find evidence that this change is because of differences in relative sensitivities of the mechanisms, and to differences in fundamental mechanism impulse responses.

Orientation sensitivity in human visual motion processing.

Orientation tuning of receptive fields is well documented in the spatial domain, but considerable variability exists amongst published estimates of orientation sensitivity of motion receptive fields. We used a two-frame motion sequence, in which one frame was binary noise and the other was a horizontally displaced and filtered version of the same noise field, to examine the orientation sensitivity of human motion mechanisms. Initially, orientations orthogonal to the direction of motion were removed from each filtered frame. Observers indicated perceived direction of motion in a single interval, binary choice task. D(max) was determined for different amounts of removed orientations, and found to remain constant across the removal of energy up to approximately +/-60 deg from vertical. In a second experiment, the orientations removed were now parallel to the direction of motion of the stimulus. D(max) fell as a cosine function with increasing removal of orientation information, in agreement with off-orientation looking or matched filtering predictions. The two experiments show the presence of mechanisms both broadly tuned and more narrowly tuned for orientation. A control experiment introduced an interstimulus interval between the two frames of our motion sequence. Performance on the direction discrimination task was severely degraded, indicating that the original results are not explicable in terms of a feature-tracking or long-range motion process. The presence of both broadly and narrowly tuned mechanisms implies multiple possible solutions to the processing of coherent plaid motion.

What does the Ternus display tell us about motion processing in human vision?

The Ternus display is a moving visual stimulus which elicits two very different percepts, according to the length of the interstimulus interval (ISI) between each frame of the motion sequence. These two percepts, referred to as element motion and group motion, have previously been analysed in terms of the operation of a low-level, dedicated short-range motion process (in the case of element motion), and of a higher-level, attentional long-range motion process (in the case of group motion). We used a novel Ternus configuration to show that both element and group motion are, in fact, mediated solely by a process sensitive to changes in the spatial appearance of the Ternus elements. In light of this, it appears that Ternus displays tell us nothing about low-level motion processing, implying that previous studies using Ternus displays, for instance those dealing with dyslexia, require reinterpretation. Further manipulations of the Ternus display revealed that the orientation and spatial-frequency discrimination of the process underlying the analysis of Ternus displays is far worse than thresholds for spatial vision. We conclude that Ternus displays are analysed via a long-range motion, or feature-tracking, process, and that this process is distinct from spatial vision.

Contour interaction for an easily resolvable stimulus.

It is well known that adjacent contours can reduce the visual acuity of single letters. Although this has traditionally been considered only in terms of a neural-based interaction, it has recently been suggested that the information content in the stimulus may account for the interaction. Here we ask the question, "Do similar interference effects occur for the discrimination of low-contrast letters whose size is larger than that corresponding to the resolution limit?" If so, previous acuity-based interaction results may be of more general importance. We show that while there are interference effects of nearby contours, they are of a form different from that observed at the resolution limit. In particular, the contrast polarity of the nearby contour is unimportant, which in turn suggests that a physical explanation is inappropriate. Also, the discrimination of an easily resolvable, unflanked Landolt C target requires information over a much wider spatial-frequency range than its counterpart at the resolution limit.

Centrifugal bias for second-order but not first-order motion.

Limited-lifetime Gabor stimuli were used to assess both first- and second-order motion in peripheral vision. Both first- and second-order motion mechanisms were present at a 20-deg eccentricity. Second-order motion, unlike first-order, exhibits a bias for centrifugal motion, suggesting a role for the second-order mechanism in optic flow processing.

Detection of constrast-defined shape.

We assessed the accuracy of contrast-defined shape detection of stimuli of constant aspect ratio, namely, circular bandpass stimuli whose radii were sinusoidally varied about a mean radius. Performance for these contrast-defined shapes, which we show is determined by the global rather than the local attributes of the stimulus, is 2-8 times worse than that for their luminance-defined counterparts, suggesting separate processing limitations. By spatially and orientationally filtering the two-dimensional fractal-noise carriers of which these stimuli were composed, we determined whether there are specific rules concerning the spatial and orientational input to shape detectors from mechanisms sensitive to the carrier structure. The results suggest that second-order circularity detectors receive mixed input from spatial-frequency-tuned and orientationally tuned cells.

Contour interaction in amblyopia: scale selection.

It has been known for some time that visual acuity in amblyopia is higher for single letters than for letters in a row (termed crowding). Early work showed that this could not be accounted for on the basis of the destructive interaction of adjacent contours (termed contour interaction), which was shown to be, in resolution units, normal in amblyopia. We have re-examined this issue using a letter stimulus that is modulated about a mean light level. This allows an examination of the effects of contrast polarity and spatial filtering within the contour interaction paradigm. We show that the majority of strabismic amblyopes that we investigated exhibit an anomalous contour interaction that, in some cases, was dependent on the contrast polarity of the flanking stimuli. Furthermore, we show that while amblyopes do select the optimum scale of analysis for unflanked stimuli, they do not select the optimum scale of analysis for flanked stimuli. For reasons that may have to do with their poorer shape discrimination, they select a non-optimal scale to process flanked stimuli.

The cortical deficit in humans with strabismic amblyopia.

To further our understanding of the cortical deficit in strabismic amblyopia, we measured, compared and mapped functional magnetic resonance imaging (fMRI) activation between the fixing and fellow amblyopic eyes of ten strabismic amblyopes. Of specific concern was whether the function of any visual area was spared in strabismic amblyopia, as recently suggested by both positron emission tomography (PET) and fMRI studies, and whether there was a close relationship between the fMRI response and known psychophysical deficits. To answer these questions we measured the psychophysical deficit in each subject and used stimuli whose relationship to the psychophysical deficit was known. We observed that stimuli that were well within the amblyopic passband did produce reduced fMRI activation, even in visual area V1. This suggests that V1 is anomalous in amblyopia. A similar level of reduction was observed in V2. In two subjects, we found that stimuli outside the amblyopic passband produced activation in visual area V3A. We did not find a close relationship between the fMRI response reduction in amblyopia and either of the known psychophysical deficits even though the fMRI response reduction in amblyopia did covary with stimulus spatial frequency.