Binocular contrast vision at and above threshold

A fundamental problem for any visual system with binocular overlap is the combination of information from the two eyes. Electrophysiology shows that binocular integration of luminance contrast occurs early in visual cortex, but a specific systems architecture has not been established for human vision. Here, we address this by performing binocular summation and monocular, binocular, and dichoptic masking experiments for horizontal 1 cycle per degree test and masking gratings. These data reject three previously published proposals, each of which predict too little binocular summation and insufficient dichoptic facilitation. However, a simple development of one of the rejected models (the twin summation model) and a completely new model (the two-stage model) provide very good fits to the data. Two features common to both models are gently accelerating (almost linear) contrast transduction prior to binocular summation and suppressive ocular interactions that contribute to contrast gain control. With all model parameters fixed, both models correctly predict (1) systematic variation in psychometric slopes, (2) dichoptic contrast matching, and (3) high levels of binocular summation for various levels of binocular pedestal contrast. A review of evidence from elsewhere leads us to favor the two-stage model. © 2006 ARVO.

Psychophysical evidence for two routes to suppression before binocular summation of signals in human vision

Visual mechanisms in primary visual cortex are suppressed by the superposition of gratings perpendicular to their preferred orientations. A clear picture of this process is needed to (i) inform functional architecture of image-processing models, (ii) identify the pathways available to support binocular rivalry, and (iii) generally advance our understanding of early vision. Here we use monoptic sine-wave gratings and cross-orientation masking (XOM) to reveal two cross-oriented suppressive pathways in humans, both of which occur before full binocular summation of signals. One is a within-eye (ipsiocular) pathway that is spatially broadband, immune to contrast adaptation and has a suppressive weight that tends to decrease with stimulus duration. The other pathway operates between the eyes (interocular), is spatially tuned, desensitizes with contrast adaptation and has a suppressive weight that increases with stimulus duration. When cross-oriented masks are presented to both eyes, masking is enhanced or diminished for conditions in which either ipsiocular or interocular pathways dominate masking, respectively. We propose that ipsiocular suppression precedes the influence of interocular suppression and tentatively associate the two effects with the lateral geniculate nucleus (or retina) and the visual cortex respectively. The interocular route is a good candidate for the initial pathway involved in binocular rivalry and predicts that interocular cross-orientation suppression should be found in cortical cells with predominantly ipsiocular drive. © 2007 IBRO.

Binocular interaction:Contrast matching and contrast discrimination are predicted by the same model

How do signals from the 2 eyes combine and interact? Our recent work has challenged earlier schemes in which monocular contrast signals are subject to square-law transduction followed by summation across eyes and binocular gain control. Much more successful was a new 'two-stage' model in which the initial transducer was almost linear and contrast gain control occurred both pre- and post-binocular summation. Here we extend that work by: (i) exploring the two-dimensional stimulus space (defined by left- and right-eye contrasts) more thoroughly, and (ii) performing contrast discrimination and contrast matching tasks for the same stimuli. Twenty-five base-stimuli made from 1 c/deg patches of horizontal grating, were defined by the factorial combination of 5 contrasts for the left eye (0.3-32%) with five contrasts for the right eye (0.3-32%). Other than in contrast, the gratings in the two eyes were identical. In a 2IFC discrimination task, the base-stimuli were masks (pedestals), where the contrast increment was presented to one eye only. In a matching task, the base-stimuli were standards to which observers matched the contrast of either a monocular or binocular test grating. In the model, discrimination depends on the local gradient of the observer's internal contrast-response function, while matching equates the magnitude (rather than gradient) of response to the test and standard. With all model parameters fixed by previous work, the two-stage model successfully predicted both the discrimination and the matching data and was much more successful than linear or quadratic binocular summation models. These results show that performance measures and perception (contrast discrimination and contrast matching) can be understood in the same theoretical framework for binocular contrast vision. © 2007 VSP.

Equivalence of physical and perceived speed in binocular rivalry

The relative dominance of gratings engaged in binocular rivalry can be influenced by their surroundings. One striking example occurs when surrounding motion is congruent with one but not the other grating (C. L. Paffen, S. F. te Pas, R. Kanai, M. J. van der Smagt, & F. A. Verstraten, 2004). However, such center-surround stimulus configurations can also modulate perceived speed, via a directionally tuned process (H. P. Norman, J. F. Norman, J. T. Todd, & D. T. Lindsey, 1996). We recorded rivalry for Gabor patches embedded in a drifting noise texture. Gratings whose directions opposed the background motion tended to dominate more, and vice versa, consistent with previous findings. Observers then matched the speed of a drifting noise-embedded Gabor to that of a Gabor surrounded by mean luminance. Surround motion produced substantial changes in perceived speed, by at least a factor of two for all observers. We then asked whether perceived speed could account for the contextual effects on dominance. We measured the effects of speed on rivalry dominance by changing the physical speeds of rivaling gratings, as determined by the matching data. We found the same pattern of dominance as for the context experiment, indicating that perceived and true speed influence rivalry in the same manner. We propose a Bayesian interpretation of the perceived speed illusion.

On the relation between dichoptic masking and binocular rivalry

When our two eyes view incompatible images, the brain invokes suppressive processes to inhibit one image, and favor the other. Two phenomena are typically observed: dichoptic masking (reduced sensitivity to one image) for brief presentations, and binocular rivalry (alternation between the two images), over longer exposures. However, it is not clear if these two phenomena arise from a common suppressive process. We investigated this by measuring both threshold elevation in simultaneous dichoptic masking and mean percept durations in rivalry, whilst varying relative stimulus orientation. Masking and rivalry showed significant correlations, such that strong masking was associated with long dominance durations. A second experiment suggested that individual differences across both measures are also correlated. These findings are consistent with varying the magnitude of interocular suppression in computational models of both rivalry and masking, and imply the existence of a common suppressive process. Since dichoptic masking has been localised to the monocular neurons of V1, this is a plausible first stage of binocular rivalry.

Binocular summation of contrast remains intact in strabismic amblyopia

PURPOSE. Strabismic amblyopia is typically associated with several visual deficits, including loss of contrast sensitivity in the amblyopic eye and abnormal binocular vision. Binocular summation ratios (BSRs) are usually assessed by comparing contrast sensitivity for binocular stimuli (sens BIN) with that measured in the good eye alone (sensGOOD), giving BSR = sensBIN/sensGOOD. This calculation provides an operational index of clinical binocular function, but does not assess whether neuronal mechanisms for binocular summation of contrast remain intact. This study was conducted to investigate this question. METHODS. Horizontal sine-wave gratings were used as stimuli (3 or 9 cyc/deg; 200 ms), and the conventional method of assessment (above) was compared with one in which the contrast in the amblyopic eye was adjusted (normalized) to equate monocular sensitivities. RESULTS. In nine strabismic amblyopes (mean age, 32 years), the results confirmed that the BSR was close to unity when the conventional method was used (little or no binocular advantage), but increased to approximately √2 or higher when the normalization method was used. The results were similar to those for normal control subjects (n = 3; mean age, 38 years) and were consistent with the physiological summation of contrast between the eyes. When the normal observers performed the experiments with a neutral-density (ND) filter in front of one eye, their performance was similar to that of the amblyopes in both methods of assessment. CONCLUSIONS. The results indicate that strabismic amblyopes have mechanisms for binocular summation of contrast and that the amblyopic deficits of binocularity can be simulated with an ND filter. The implications of these results for best clinical practice are discussed. Copyright © Association for Research in Vision and Ophthalmology.

Natural images dominate in binocular rivalry

Ecological approaches to perception have demonstrated that information encoding by the visual system is informed by the natural environment, both in terms of simple image attributes like luminance and contrast, and more complex relationships corresponding to Gestalt principles of perceptual organization. Here, we ask if this optimization biases perception of visual inputs that are perceptually bistable. Using the binocular rivalry paradigm, we designed stimuli that varied in either their spatiotemporal amplitude spectra or their phase spectra. We found that noise stimuli with “natural” amplitude spectra (i.e., amplitude content proportional to 1/f, where f is spatial or temporal frequency) dominate over those with any other systematic spectral slope, along both spatial and temporal dimensions. This could not be explained by perceived contrast measurements, and occurred even though all stimuli had equal energy. Calculating the effective contrast following attenuation by a model contrast sensitivity function suggested that the strong contrast dependency of rivalry provides the mechanism by which binocular vision is optimized for viewing natural images. We also compared rivalry between natural and phase-scrambled images and found a strong preference for natural phase spectra that could not be accounted for by observer biases in a control task. We propose that this phase specificity relates to contour information, and arises either from the activity of V1 complex cells, or from later visual areas, consistent with recent neuroimaging and single-cell work. Our findings demonstrate that human vision integrates information across space, time, and phase to select the input most likely to hold behavioral relevance.

Nonlinearities in the binocular combination of luminance and contrast

We studied the rules by which visual responses to luminous targets are combined across the two eyes. Previous work has found very different forms of binocular combination for targets defined by increments and by decrements of luminance, with decrement data implying a severe nonlinearity before binocular combination. We ask whether this difference is due to the luminance of the target, the luminance of the background, or the sign of the luminance excursion. We estimated the pre-binocular nonlinearity (power exponent) by fitting a computational model to ocular equibrightness matches. The severity of the nonlinearity had a monotonic dependence on the signed difference between target and background luminance. For dual targets, in which there was both a luminance increment and a luminance decrement (e.g. contrast), perception was governed largely by the decrement. The asymmetry in the nonlinearities derived from the subjective matching data made a clear prediction for visual performance: there should be more binocular summation for detecting luminance increments than for detecting luminance decrements. This prediction was confirmed by the results of a subsequent experiment. We discuss the relation between these results and luminance nonlinearities such as a logarithmic transform, as well as the involvement of contemporary model architectures of binocular vision.

Binocular combination at threshold: temporal filtering and summation of signals in separate ON and OFF channels

How does the brain combine spatio-temporal signals from the two eyes? We quantified binocular summation as the improvement in 2AFC contrast sensitivity for flickering gratings seen by two eyes compared with one. Binocular gratings in-phase showed sensitivity up to 1.8 times higher, suggesting nearly linear summation of contrasts. The binocular advantage decreased to 1.4 at lower spatial and higher temporal frequencies (0.25 cycle deg-1, 30 Hz). Dichoptic, antiphase gratings showed only a small binocular advantage, by a factor of 1.1 to 1.2, but no evidence of cancellation. We present a signal-processing model to account for the contrast-sensitivity functions and the pattern of binocular summation. It has linear sustained and transient temporal filters, nonlinear transduction, and half-wave rectification that creates ON and OFF channels. Binocular summation occurs separately within ON and OFF channels, thus explaining the phase-specific binocular advantage. The model also accounts for earlier findings on detection of brief antiphase flashes and the surprising finding that dichoptic antiphase flicker is seen as frequency-doubled (Cavonius et al, 1992 Ophthalmic and Physiological Optics 12 153 - 156). [Supported by EPSRC project GR/S74515/01].

Binocular summation at contrast threshold:a new look

Contrast sensitivity is better with two eyes than one. The standard view is that thresholds are about 1.4 (v2) times better with two eyes, and that this arises from monocular responses that, near threshold, are proportional to the square of contrast, followed by binocular summation of the two monocular signals. However, estimates of the threshold ratio in the literature vary from about 1.2 to 1.9, and many early studies had methodological weaknesses. We collected extensive new data, and applied a general model of binocular summation to interpret the threshold ratio. We used horizontal gratings (0.25 - 4 cycles deg-1) flickering sinusoidally (1 - 16 Hz), presented to one or both eyes through frame-alternating ferroelectric goggles with negligible cross-talk, and used a 2AFC staircase method to estimate contrast thresholds and psychometric slopes. Four naive observers completed 20 000 trials each, and their mean threshold ratios were 1.63, 1.69, 1.71, 1.81 - grand mean 1.71 - well above the classical v2. Mean ratios tended to be slightly lower (~1.60) at low spatial or high temporal frequencies. We modelled contrast detection very simply by assuming a single binocular mechanism whose response is proportional to (Lm + Rm) p, followed by fixed additive noise, where L,R are contrasts in the left and right eyes, and m, p are constants. Contrast-gain-control effects were assumed to be negligible near threshold. On this model the threshold ratio is 2(?1/m), implying that m=1.3 on average, while the Weibull psychometric slope (median 3.28) equals 1.247mp, yielding p=2.0. Together, the model and data suggest that, at low contrasts across a wide spatiotemporal frequency range, monocular pathways are nearly linear in their contrast response (m close to 1), while a strongly accelerating nonlinearity (p=2, a 'soft threshold') occurs after binocular summation. [Supported by EPSRC project grant GR/S74515/01]

Contrast masking in strabismic amblyopia: attenuation, noise, interocular suppression and binocular summation

To investigate amblyopic contrast vision at threshold and above we performed pedestal-masking (contrastdiscrimination) experiments with a group of eight strabismic amblyopes using horizontal sinusoidal gratings (mainly 3 c/deg) in monocular, binocular and dichoptic configurations balanced across eye (i.e. five conditions). With some exceptions in some observers, the four main results were as follows. (1) For the monocular and dichoptic conditions, sensitivity was less in the amblyopic eye than in the good eye at all mask contrasts. (2) Binocular and monocular dipper functions superimposed in the good eye. (3) Monocular masking functions had a normal dipper shape in the good eye, but facilitation was diminished in the amblyopic eye. (4) A less consistent result was normal facilitation in dichoptic masking when testing the good eye, but a loss of this when testing the amblyopic eye. This pattern of amblyopic results was replicated in a normal observer by placing a neutral density filter in front of one eye. The two-stage model of binocular contrast gain control [Meese, T.S., Georgeson, M.A. & Baker, D.H. (2006). Binocular contrast vision at and above threshold. Journal of Vision 6, 1224--1243.] was `lesioned' in several ways to assess the form of the amblyopic deficit. The most successful model involves attenuation of signal and an increase in noise in the amblyopic eye, and intact stages of interocular suppression and binocular summation. This implies a behavioural influence from monocular noise in the amblyopic visual system as well as in normal observers with an ND filter over one eye.

Binocular contrast interactions:Dichoptic masking is not a single process

To decouple interocular suppression and binocular summation we varied the relative phase of mask and target in a 2IFC contrast-masking paradigm. In Experiment I, dichoptic mask gratings had the same orientation and spatial frequency as the target. For in-phase masking, suppression was strong (a log-log slope of ∼1) and there was weak facilitation at low mask contrasts. Anti-phase masking was weaker (a log-log slope of ∼0.7) and there was no facilitation. A two-stage model of contrast gain control [Meese, T.S., Georgeson, M.A. and Baker, D.H. (2006). Binocular contrast vision at and above threshold. Journal of Vision, 6: 1224-1243] provided a good fit to the in-phase results and fixed its free parameters. It made successful predictions (with no free parameters) for the anti-phase results when (A) interocular suppression was phase-indifferent but (B) binocular summation was phase sensitive. Experiments II and III showed that interocular suppression comprised two components: (i) a tuned effect with an orientation bandwidth of ∼±33° and a spatial frequency bandwidth of >3 octaves, and (ii) an untuned effect that elevated threshold by a factor of between 2 and 4. Operationally, binocular summation was more tightly tuned, having an orientation bandwidth of ∼±8°, and a spatial frequency bandwidth of ∼0.5 octaves. Our results replicate the unusual shapes of the in-phase dichoptic tuning functions reported by Legge [Legge, G.E. (1979). Spatial frequency masking in human vision: Binocular interactions. Journal of the Optical Society of America, 69: 838-847]. These can now be seen as the envelope of the direct effects from interocular suppression and the indirect effect from binocular summation, which contaminates the signal channel with a mask that has been suppressed by the target. © 2007 Elsevier Ltd. All rights reserved.

Neuronal convergence in early contrast vision: binocular summation is followed by response nonlinearity and area summation

We assessed summation of contrast across eyes and area at detection threshold ( C t). Stimuli were sine-wave gratings (2.5 c/deg) spatially modulated by cosine- and anticosine-phase raised plaids (0.5 c/deg components oriented at ±45°). When presented dichoptically the signal regions were interdigitated across eyes but produced a smooth continuous grating following their linear binocular sum. The average summation ratio ( C t1/([ C t1+2]) for this stimulus pair was 1.64 (4.3 dB). This was only slightly less than the binocular summation found for the same patch type presented to both eyes, and the area summation found for the two different patch types presented to the same eye. We considered 192 model architectures containing each of the following four elements in all possible orders: (i) linear summation or a MAX operator across eyes, (ii) linear summation or a MAX operator across area, (iii) linear or accelerating contrast transduction, and (iv) additive Gaussian, stochastic noise. Formal equivalences reduced this to 62 different models. The most successful four-element model was: linear summation across eyes followed by nonlinear contrast transduction, linear summation across area, and late noise. Model performance was enhanced when additional nonlinearities were placed before binocular summation and after area summation. The implications for models of probability summation and uncertainty are discussed.

Visual consciousness: the binocular rivalry explosion

A new behavioural technique solves a long-standing puzzle of binocular suppression, demonstrating that adapting reciprocal inhibition governs visual sensitivity, and raising key questions about visual awareness.

Binocular fusion, suppression and diplopia for blurred edges

Purpose: (1) To devise a model-based method for estimating the probabilities of binocular fusion, interocular suppression and diplopia from psychophysical judgements, (2) To map out the way fusion, suppression and diplopia vary with binocular disparity and blur of single edges shown to each eye, (3) To compare the binocular interactions found for edges of the same vs opposite contrast polarity. Methods: Test images were single, horizontal, Gaussian-blurred edges, with blur B = 1-32 min arc, and vertical disparity 0-8.B, shown for 200 ms. In the main experiment, observers reported whether they saw one central edge, one offset edge, or two edges. We argue that the relation between these three response categories and the three perceptual states (fusion, suppression, diplopia) is indirect and likely to be distorted by positional noise and criterion effects, and so we developed a descriptive, probabilistic model to estimate both the perceptual states and the noise/criterion parameters from the data. Results: (1) Using simulated data, we validated the model-based method by showing that it recovered fairly accurately the disparity ranges for fusion and suppression, (2) The disparity range for fusion (Panum's limit) increased greatly with blur, in line with previous studies. The disparity range for suppression was similar to the fusion limit at large blurs, but two or three times the fusion limit at small blurs. This meant that diplopia was much more prevalent at larger blurs, (3) Diplopia was much more frequent when the two edges had opposite contrast polarity. A formal comparison of models indicated that fusion occurs for same, but not opposite, polarities. Probability of suppression was greater for unequal contrasts, and it was always the lower-contrast edge that was suppressed. Conclusions: Our model-based data analysis offers a useful tool for probing binocular fusion and suppression psychophysically. The disparity range for fusion increased with edge blur but fell short of complete scale-invariance. The disparity range for suppression also increased with blur but was not close to scale-invariance. Single vision occurs through fusion, but also beyond the fusion range, through suppression. Thus suppression can serve as a mechanism for extending single vision to larger disparities, but mainly for sharper edges where the fusion range is small (5-10 min arc). For large blurs the fusion range is so much larger that no such extension may be needed. © 2014 The College of Optometrists.