Gain control of saccadic eye movements is probabilistic.

Saccades are rapid eye movements that orient the visual axis toward objects of interest to allow their processing by the central, high-acuity retina. Our ability to collect visual information efficiently relies on saccadic accuracy, which is limited by a combination of uncertainty in the location of the target and motor noise. It has been observed that saccades have a systematic tendency to fall short of their intended targets, and it has been suggested that this bias originates from a cost function that overly penalizes hypermetric errors. Here, we tested this hypothesis by systematically manipulating the positional uncertainty of saccadic targets. We found that increasing uncertainty produced not only a larger spread of the saccadic endpoints but also more hypometric errors and a systematic bias toward the average of target locations in a given block, revealing that prior knowledge was integrated into saccadic planning. Moreover, by examining how variability and bias covaried across conditions, we estimated the asymmetry of the cost function and found that it was related to individual differences in the additional time needed to program secondary saccades for correcting hypermetric errors, relative to hypometric ones. Taken together, these findings reveal that the saccadic system uses a probabilistic-Bayesian control strategy to compensate for uncertainty in a statistically principled way and to minimize the expected cost of saccadic errors.

Models for discriminating image blur from loss of contrast.

Observers can discriminate between blurry and low-contrast images (Morgan, 2017). Wang and Simoncelli (2004) demonstrated that a code for blur is inherent to the phase relationships between localized pattern detectors of different scales. To test whether human observers actually use local phase coherence when discriminating between image blur and loss of contrast, we compared phase-scrambled chessboards with unscrambled chessboards. Although both stimuli had identical amplitude spectra, local phase coherence was disrupted by phase-scrambling. Human observers were required to concurrently detect and identify (as contrast or blur) image manipulations in the 2 × 2 forced-choice paradigm (Nachmias & Weber, 1975; Watson & Robson, 1981) traditionally considered to be a litmus test for "labelled lines" (i.e. detection mechanisms that can be distinguished on the basis of their preferred stimuli). Phase scrambling reduced some observers' ability to discriminate between blur and a reduction in contrast. However, none of our observers produced data consistent with Watson and Robson's most stringent test for labeled lines, regardless whether phases were scrambled or not. Models of performance fit significantly better when (a) the blur detector also responded to contrast modulations, (b) the contrast detector also responded to blur modulations, or (c) noise in the two detectors was anticorrelated.

A perceptual bias for man-made objects in humans.

Ambiguous images are widely recognized as a valuable tool for probing human perception. Perceptual biases that arise when people make judgements about ambiguous images reveal their expectations about the environment. While perceptual biases in early visual processing have been well established, their existence in higher-level vision has been explored only for faces, which may be processed differently from other objects. Here we developed a new, highly versatile method of creating ambiguous hybrid images comprising two component objects belonging to distinct categories. We used these hybrids to measure perceptual biases in object classification and found that images of man-made (manufactured) objects dominated those of naturally occurring (non-man-made) ones in hybrids. This dominance generalized to a broad range of object categories, persisted when the horizontal and vertical elements that dominate man-made objects were removed and increased with the real-world size of the manufactured object. Our findings show for the first time that people have perceptual biases to see man-made objects and suggest that extended exposure to manufactured environments in our urban-living participants has changed the way that they see the world.

Calculation Efficiencies for Mean Numerosity.

Relative numerosity is traditionally studied using texture pairs. Observers must decide which member of each pair has the greater total number of texture elements. In the present experiment, textures were segregated into nonoverlapping "sectors" containing between zero and four elements, and our observers were asked to select the texture containing the greater average number of texture elements (per sector). If observers were more sensitive to total numerosity than average numerosity, their performance (quantified by the just-noticeable Weber fraction) should have been better when the two textures occupied the same number of sectors than when they occupied unequal numbers of sectors. However, we recorded Weber fractions between 11% and 13% for all observers in all conditions. This performance was comparable with an otherwise-ideal observer whose decisions were based on between three and five sectors in each texture. We conjecture that traditional numerosity discriminations are based on similarly small numbers of element clusters.

Robust averaging protects decisions from noise in neural computations.

An ideal observer will give equivalent weight to sources of information that are equally reliable. However, when averaging visual information, human observers tend to downweight or discount features that are relatively outlying or deviant ('robust averaging'). Why humans adopt an integration policy that discards important decision information remains unknown. Here, observers were asked to judge the average tilt in a circular array of high-contrast gratings, relative to an orientation boundary defined by a central reference grating. Observers showed robust averaging of orientation, but the extent to which they did so was a positive predictor of their overall performance. Using computational simulations, we show that although robust averaging is suboptimal for a perfect integrator, it paradoxically enhances performance in the presence of "late" noise, i.e. which corrupts decisions during integration. In other words, robust decision strategies increase the brain's resilience to noise arising in neural computations during decision-making.

Improvement of contrast sensitivity with practice is not compatible with a sensory threshold account.

In forced-choice detection, incorrect responses are routinely ascribed to internal noise, because experienced psychophysical observers do not act as if they have a sensory threshold, below which all perceived intensities would be identical. To determine whether inexperienced observers have sensory thresholds, we examined psychometric functions (percent correct versus log contrast) for detection and detection in full-screen, dynamic visual noise. Over five days, neither type of psychometric function changed shape, but both shifted leftwards, indicating increased sensitivity. These results are not consistent with a lowered sensory threshold, which would decrease psychometric slope. Our results can be understood within the context of Dosher and Lu's "stochastic" perceptual template model [Vis. Res.40, 1269 (2000)], augmented to allow intrinsic uncertainty. Specifically, our results are consistent with a combination of reduced internal additive noise and improved filtering of external noise.

Inefficiency of orientation averaging: Evidence for hybrid serial/parallel temporal integration.

Intuition suggests that increased viewing time should allow for the accumulation of more visual information, but scant support for this idea has been found in studies of voluntary averaging, where observers are asked to make decisions based on perceived average size. In this paper we examine the dynamics of information accrual in an orientation-averaging task. With orientation (unlike intensive dimensions such as size), it is relatively safe to use an item's physical value as an approximation for its average perceived value. We displayed arrays containing eight iso-eccentric Gabor patterns, and asked six trained psychophysical observers to compare their average orientation with that of probe stimuli that were visible before, during, or only after the presentation of the Gabor array. From the relationship between orientation variance and human performance, we obtained estimates of effective set size, i.e., the number of items that an ideal observer would need to assess in order to estimate average orientation as well as our human observers did. We found that display duration had only a modest influence on effective set size. It rose from an average of ∼2 for 0.1-s displays to an average of ∼3 for 3.3-s displays. These results suggest that the visual computation is neither purely serial nor purely parallel. Computations of this nature can be made with a hybrid process that takes a series of subsamples of a few elements at a time.

Tilted frames of reference have similar effects on the perception of gravitational vertical and the planning of vertical saccadic eye movements.

We investigated the effects of a tilted reference frame (i.e., allocentric visual context) on the perception of the gravitational vertical and saccadic eye movements along a planned egocentric vertical path. Participants (n = 5) in a darkened room fixated a point in the center of a circle on an LCD display and decided which of two sequentially presented dots was closer to the unmarked '6 o'clock' position on that circle (i.e., straight down toward their feet). The slope of their perceptual psychometric functions showed that participants were able to locate which dot was nearer the vertical with a precision of 1°-2°. For three of the participants, a square frame centered at fixation and tilted (in the roll direction) 5.6° from the vertical caused a strong perceptual bias, manifest as a shift in the psychometric function, in the direction of the traditional 'rod-and-frame' effect, without affecting precision. The other two participants showed negligible or no equivalent biases. The same subjects participated in the saccade version of the task, in which they were instructed to shift their gaze to the 6 o'clock position as soon as the central fixation point disappeared. The participants who showed perceptual biases showed biases of similar magnitude in their saccadic endpoints, with a strong correlation between perceptual and saccadic biases across all subjects. Tilting of the head 5.6° reduced both perceptual and saccadic biases in all but one observer, who developed a strong saccadic bias. Otherwise, the overall pattern and significant correlations between results remained the same. We conclude that our observers' saccades-to-vertical were dominated by perceptual input, which outweighed any gravitational or head-centered input.

Connecting psychophysical performance to neuronal response properties I: Discrimination of suprathreshold stimuli.

One of the major goals of sensory neuroscience is to understand how an organism's perceptual abilities relate to the underlying physiology. To this end, we derived equations to estimate the best possible psychophysical discrimination performance, given the properties of the neurons carrying the sensory code.We set up a generic sensory coding model with neurons characterized by their tuning function to the stimulus and the random process that generates spikes. The tuning function was a Gaussian function or a sigmoid (Naka-Rushton) function.Spikes were generated using Poisson spiking processes whose rates were modulated by a multiplicative, gamma-distributed gain signal that was shared between neurons. This doubly stochastic process generates realistic levels of neuronal variability and a realistic correlation structure within the population. Using Fisher information as a close approximation of the model's decoding precision, we derived equations to predict the model's discrimination performance from the neuronal parameters. We then verified the accuracy of our equations using Monte Carlo simulations. Our work has two major benefits. Firstly, we can quickly calculate the performance of physiologically plausible population-coding models by evaluating simple equations, which makes it easy to fit the model to psychophysical data. Secondly, the equations revealed some remarkably straightforward relationships between psychophysical discrimination performance and the parameters of the neuronal population, giving deep insights into the relationships between an organism's perceptual abilities and the properties of the neurons on which those abilities depend.

Connecting psychophysical performance to neuronal response properties II: Contrast decoding and detection.

The purpose of this article is to provide mathematical insights into the results of some Monte Carlo simulations published by Tolhurst and colleagues (Clatworthy, Chirimuuta, Lauritzen, & Tolhurst, 2003; Chirimuuta & Tolhurst, 2005a). In these simulations, the contrast of a visual stimulus was encoded by a model spiking neuron or a set of such neurons. The mean spike count of each neuron was given by a sigmoidal function of contrast, the Naka-Rushton function. The actual number of spikes generated on each trial was determined by a doubly stochastic Poisson process. The spike counts were decoded using a Bayesian decoder to give an estimate of the stimulus contrast. Tolhurst and colleagues used the estimated contrast values to assess the model's performance in a number of ways, and they uncovered several relationships between properties of the neurons and characteristics of performance. Although this work made a substantial contribution to our understanding of the links between physiology and perceptual performance, the Monte Carlo simulations provided little insight into why the obtained patterns of results arose or how general they are. We overcame these problems by deriving equations that predict the model's performance. We derived an approximation of the model's decoding precision using Fisher information. We also analyzed the model's contrast detection performance and discovered a previously unknown theoretical connection between the Naka-Rushton contrast-response function and the Weibull psychometric function. Our equations give many insights into the theoretical relationships between physiology and perceptual performance reported by Tolhurst and colleagues, explaining how they arise and how they generalize across the neuronal parameter space.

Awareness is the key to attraction: dissociating the tilt illusions via conscious perception.

The tilt illusion is a compelling example of contextual influence exerted by an oriented surround on a target's perceived orientation. A vertical target appears to be tilted away from a 15° oriented surround but appears to be tilted toward a 75° tilted surround. We tested the claim that these biases result from distinct sensory processes: a low-level repulsive process and a higher-level attractive process. If this claim were correct, then surround visibility would be a requirement for attraction, but it would not necessarily be a requirement for repulsion. Indeed, Motoyoshi and Hayakawa (2010) have demonstrated that repulsion can survive removal of the surround from phenomenal awareness using adaptation-induced blindness. Here we sought to test this prediction by measuring the orientation biases in a parafoveally presented Gabor patch surrounded by tilted gratings after 20-s adaptation. The adapting stimulus was an annularly windowed plaid composed of vertical and horizontal jittering gratings. Observers were instructed to maintain fixation throughout the trial and report whether the Gabor appeared to be tilted clockwise or anticlockwise of vertical. They also had to indicate whether the surround was visible after adaptation. Postadaptation biases were then compared with those obtained in a control experiment without dynamic adaptation. We found large repulsive biases induced by 15° oriented surrounds, but no attractive biases were induced by 75° tilted surrounds. This result shows that attractive effects do require visual awareness and thereby provides robust evidence for the existence of two separate mechanisms mediating the phenomenology of the tilt illusions.

Summary statistics for size over space and time.

A number of studies have investigated how the visual system extracts the average feature-value of an ensemble of simultaneously or sequentially delivered stimuli. In this study we model these two processes within the unitary framework of linear systems theory. The specific feature value used in this investigation is size, which we define as the logarithm of a circle's diameter. Within each ensemble, sizes were drawn from a normal distribution. Average size discrimination was measured using ensembles of one and eight circles. These circles were presented simultaneously (display times: 13-427 ms), one at a time, or eight at a time (temporal-frequencies: 1.2-38 Hz). Thresholds for eight-item ensembles were lower than thresholds for one-item ensembles. Thresholds decreased by a factor of 1.3 for a 3,200% increase in display time, and decreased by the same factor for a 3,200% decrease in temporal frequency. Modeling and simulations show that the data are consistent with one readout of three to four items every 210 ms.

Spatial summation for motion detection.

We used the psychophysical summation paradigm to reveal some spatial characteristics of the mechanism responsible for detecting a motion-defined visual target in central vision. There has been much previous work on spatial summation for motion detection and direction discrimination, but none has assessed it in terms of the velocity threshold or used velocity noise to provide a measure of the efficiency of the velocity processing mechanism. Motion-defined targets were centered within square fields of randomly selected gray levels. The motion was produced within the disk-shaped target region by shifting the pixels rightwards for 0.2 s. The uniform target motion was perturbed by Gaussian motion noise in horizontal strips of 16 pixels. Independent variables were field size, the diameter of the disk target, and the variance of an independent perturbation added to the (signed) velocity of each 16-pixel strip. The dependent variable was the threshold velocity for target detection. Velocity thresholds formed swoosh-shaped (descending, then ascending) functions of target diameter. Minimum values were obtained when targets subtended approximately 2 degrees of visual angle. The data were fit with a continuum of models, extending from the theoretically ideal observer through various inefficient and noisy refinements thereof. In particular, we introduce the concept of sparse sampling to account for the relative inefficiency of the velocity thresholds. The best fits were obtained from a model observer whose responses were determined by comparing the velocity profile of each stimulus with a limited set of sparsely sampled "DoG" templates, each of which is the product of a random binary array and the difference between two 2-D Gaussian density functions.

Monocular and binocular mechanisms detect modulations of dot density and dot contrast.

Strong reciprocity has been demonstrated between (1) spatial modulations of dot density and modulations of dot luminance, and (2) modulations of dot density and modulations of dot contrast, in textures. The latter are much easier to detect when presented in phase with one another than when presented 180° out of phase, although out-of-phase modulations can also be detected given sufficient amplitude. This result supports the existence of two detection mechanisms: one that is excited by both density modulations and contrast modulations (quiescent when those modulations are presented 180° out of phase) and another that is relatively insensitive to either density modulations or contrast modulations (thus remaining stimulated regardless of phase angle). We investigate whether the mechanism responsible for detecting out-of-phase modulations depends on high-level computations (downstream from the confluence of monocular signals) or whether both mechanisms are situated at the monocular level of visual processing. Specifically, density-modulated and/or contrast-modulated stimuli were presented monocularly (i.e., to the same eye) or dichoptically (i.e., to opposite eyes). Out-of-phase modulations of density were much easier to detect when presented dichoptically. A dichoptic advantage was also found for out-of-phase density and contrast modulations. These dichoptic advantages imply conscious access to a mechanism at the monocular level of processing. When density modulations were presented dichoptically, 180° out of phase, detection thresholds were highest. Consequently, a mechanism with binocular input must also contribute to the detection of these modulations. We describe a minimal, image-based model for these results that contains one monocular computation and one binocular computation.

Comparison of Depth-Related Visuomotor Task Performance in Uniocular Individuals and in Binocular Controls With and Without Temporary Monocular Occlusion.

Purpose: Do one-eyed (uniocular) humans use monocular depth cues differently from those with intact binocularity to perform depth-related visuomotor tasks that emulate complex activities of daily living? If so, does performance depend on the participant's age, duration of uniocularity and head movements? Methods: Forty-five uniocular cases (age range 6-37 years; 2.4 months-31.0 years of uniocularity) and 46 age-similar binocular controls performed a task that required them to pass a hoop around an electrified wire convoluted in depth multiple times, while avoiding contact as indicated by auditory feedback. The task was performed with and without head restraint, in random order. The error rate and speed were calculated from the frequency of contact between the hoop and wire and the total task duration (adjusting for error time), respectively, all determined from video recordings of the task. Head movements were analyzed from the videos using face-tracking software. Results: Error rate decreased with age (P < 0.001) until the late teen years while speed revealed no such trend. Across all ages, the error rate increased and speed decreased in the absence of binocularity (P < 0.001). There was no additional error reduction with duration of uniocularity (P = 0.16). Head movements provided no advantage to task performance, despite generating parallax disparities comparable to binocular viewing. Conclusions: Performance in a dynamic, depth-related visuomotor task is reduced in the absence of binocular viewing, independent of age-related performance level. This study finds no evidence for a prolonged experience with monocular depth cues being advantageous for such tasks over transient loss of binocularity.

Aftereffect of perceived regularity.

Regularity is a ubiquitous feature of the visual world. We demonstrate that regularity is an adaptable visual dimension: The perceived regularity of a pattern is reduced following adaptation to a pattern with a similar or greater degree of regularity. Stimuli consisted of 7×7 element arrays arranged on square grids presented in a circular aperture. The position of each element was randomly jittered from its baseline position by an amount that determined its degree of irregularity. The elements of the pattern consisted of dark Gaussian blobs (GBs), difference of Gaussians (DOGs), or random binary patterns (RBPs). Observers adapted for 60 s to either a single pattern or a pair of patterns with particular regularities, and the perceived regularities of subsequently presented test patterns were measured using a conventional staircase matching procedure. We found that the regularity aftereffect (RAE) was unidirectional: Adaptation only caused test patterns to appear less regular. We also found that RAEs transferred from GB adaptors to both DOG and RBP test patterns and from DOG and RBP adaptors to GB patterns. We suggest that regularity is coded by the peakedness in the distribution of spatial-frequency channel responses across scale, and that the RAE is a result of a flattening of this distribution by adaptation. Thus, the RAE may be a consequence of contrast normalization, and an example of norm-based coding where irregularity is the norm.

The best fitting of three contemporary observer models reveals how participants' strategy influences the window of subjective synchrony.

When experimenters vary the timing between two intersensory events, and participants judge their simultaneity, an inverse-U-shaped psychometric function is obtained. Typically, this simultaneity function is first fitted with a model for each participant separately, before best-fitting parameters are utilized (e.g., compared across conditions) in the second stage of a two-step inferential procedure. Often, simultaneity-function width is interpreted as representing sensitivity to asynchrony, and/or ascribed theoretical equivalence to a window of multisensory temporal binding. Here, we instead fit a single (principled) multilevel model to data from the entire group and across several conditions at once. By asking 20 participants to sometimes be more conservative in their judgments, we demonstrate how the width of the simultaneity function is prone to strategic change and thus questionable as a measure of either sensitivity to asynchrony or multisensory binding. By repeating our analysis with three different models (two implying a decision based directly on subjective asynchrony, and a third deriving this decision from the correlation between filtered responses to sensory inputs) we find that the first model, which hypothesizes, in particular, Gaussian latency noise and difficulty maintaining the stability of decision criteria across trials, is most plausible for these data. (PsycInfo Database Record (c) 2023 APA, all rights reserved).

Recognition criteria vary with fluctuating uncertainty.

In distinct experiments we examined memories for orientation and size. After viewing a randomly oriented Gabor patch (or a plain white disk of random size), observers were given unlimited time to reproduce as faithfully as possible the orientation (or size) of that standard stimulus with an adjustable Gabor patch (or disk). Then, with this match stimulus still in view, a recognition probe was presented. On half the trials, this probe was identical to the standard. We expected observers to classify the probe (a same/different task) on the basis of its difference from the match, which should have served as an explicit memory of the standard. Observers did better than that. Larger differences were classified as "same" when probe and standard were indeed identical. In some cases, recognition performance exceeded that of a simulated observer subject to the same matching errors, but forced to adopt the single most advantageous criterion difference between the probe and match. Recognition must have used information that was not or could not be exploited in the reproduction phase. One possible source for that information is observers' confidence in their reproduction (e.g., in their memory of the standard). Simulations confirm the enhancement of recognition performance when decision criteria are adjusted trial-by-trial, on the basis of the observer's estimated reproduction error.

An image-driven model for pattern detection, resistant to Birdsall linearisation.

If detection were governed by an isolated (and possibly nonlinear) transducer, then a linearisation of the psychometric function (d-prime vs target amplitude) must accompany any threshold elevation due to the addition of external noise. This is the Birdsall theorem. From the fact that noise can elevate threshold without linearising the psychometric function, we can safely infer that detection is not governed by an isolated transducer. Heretofore, image-driven models, which accept images or numerical descriptions thereof as input, have proven incompatible with this failure of Birdsall linearisation, unless they incorporate the principle of intrinsic uncertainty, which asserts that detection is governed by the maximum activity in several independent (noisy) sensors. One image-driven model incompatible with the failure of Birdsall linearisation is Watson and Solomon's (J. Opt. Soc. Am. A, 14 (1997), 2379) model of visual contrast gain control and pattern masking. Here I report a simple modification - pooling sensor outputs before, instead of after the comparison of input images - allowing that model to predict curved psychometric functions, even when external noise elevates threshold by more than 20 dB, without any detrimental effect to the quality of its fit to pattern-masking thresholds in the absence of noise. The failure of Birdsall linearisation, therefore, does not necessarily imply independent samples of performance-limiting noise in multiple visual sensors. Instead, performance-limiting noise may arise after the visual system combines output from mutually inhibitory sensors.

Perceptual decisions and oculomotor responses rely on temporally distinct streams of evidence.

Perceptual decisions often require the integration of noisy sensory evidence over time. This process is formalized with sequential sampling models, where evidence is accumulated up to a decision threshold before a choice is made. Although intuition suggests that decision formation must precede the preparation of a motor response (i.e., the action used to communicate the choice), neurophysiological findings have suggested that these two processes might be one and the same. To test this idea, we developed a reverse-correlation protocol in which the visual stimuli that influence decisions can be distinguished from those guiding motor responses. In three experiments, we found that the temporal weighting function of oculomotor responses did not overlap with the relatively early weighting function of stimulus properties having an impact on decision formation. These results support a timeline in which perceptual decisions are formed, at least in part, prior to the preparation of a motor response.