Motion direction is represented as a bimodal probability distribution in the human visual cortex.

Humans infer motion direction from noisy sensory signals. We hypothesize that to make these inferences more precise, the visual system computes motion direction not only from velocity but also spatial orientation signals - a 'streak' created by moving objects. We implement this hypothesis in a Bayesian model, which quantifies knowledge with probability distributions, and test its predictions using psychophysics and fMRI. Using a probabilistic pattern-based analysis, we decode probability distributions of motion direction from trial-by-trial activity in the human visual cortex. Corroborating the predictions, the decoded distributions have a bimodal shape, with peaks that predict the direction and magnitude of behavioral errors. Interestingly, we observe similar bimodality in the distribution of the observers' behavioral responses across trials. Together, these results suggest that observers use spatial orientation signals when estimating motion direction. More broadly, our findings indicate that the cortical representation of low-level visual features, such as motion direction, can reflect a combination of several qualitatively distinct signals.

Perception of length and orientation in dry immersion.

Introduction: How does gravity (or lack thereof) affect sensory-motor processing? We analyze sensorimotor estimation dynamics for line segments with varying direction (orientation) in a 7-day dry immersion (DI), a ground-based model of gravitational unloading. Methods: The measurements were carried out before the start of the DI, on the first, third, fifth and seventh days of the DI, and after its completion. At the memorization stage, the volunteers led the leading hand along the visible segment on a touchscreen display, and at the reproduction stage they repeated this movement on an empty screen. A control group followed the same procedure without DI. Results: Both in the DI and control groups, when memorizing, the overall error in estimating the lengths and directions of the segments was small and did not have pronounced dynamics; when reproducing, an oblique effect (higher variability of responses to oblique orientations compared to cardinal ones) was obtained. We then separated biases (systematic error) and uncertainty (random error) in subjects' responses. At the same time, two opposite trends were more pronounced in the DI group during the DI. On the one hand the cardinal bias (a repulsion of orientation estimates away from cardinal axes) and, to a small extent, the variability of direction estimates decreased. On the other hand, the overestimation bias in length estimates increased. Discussion: Such error pattern strongly supports the hypotheses of the vector encoding, in which the direction and length of the planned movement are encoded independently of each other when the DI disrupts primarily the movement length encoding.

The influence of the tested item on serial dependence in perceptual decisions.

Serial dependence in vision reflects how perceptual decisions can be biased by what we have recently perceived. Serial dependence studies test single items' effects on perceptual decisions. However, our visual world contains multiple objects at any given moment, so it's important to understand how past experiences affect not only a single object but also perception in a more general sense. Here we asked the question: What effect does a single item have when there is more than one subsequently presented test item? We displayed a single line (inducer) at the screen center, then either a single test-line or two simultaneous test-lines, varying in orientation space to the inducer. Next, participants reported test-line orientation using a left or right located response circle (to indicate which test-line should be reported). The results demonstrated that the inducer influenced subsequent perceptual judgments of a test-line even when two test-lines were presented. Distant items caused repulsive serial dependence, while close items caused attractive serial dependence. This shows how a single inducer can influence test-line judgments, even when two test-lines are presented, and can produce attractive and repulsive serial dependence biases when the item to report is revealed after it has disappeared.

Extracting statistical information about shapes in the visual environment.

It is well known that observers can use so-called summary statistics of visual ensembles to simplify perceptual processing. The assumption has been that instead of representing feature distributions in detail the visual system extracts the mean and variance of visual ensembles. But recent evidence from implicit testing using a method called feature distribution learning showed that far more detail of the distributions is retained than the summary statistic literature indicates. Observers also encode higher-order statistics such as the kurtosis of feature distributions of orientation and color. But this sort of learning has not been shown for more intricate aspects of visual information. Here we tested the learning of distractor ensembles for shape, using the feature distribution learning method. Using a linearized circular shape space, we found that learning of detailed distributions of shape does not occur for this shape space while observers were able to learn the mean and range of the distributions. Previous demonstrations of feature distribution learning involved simpler feature dimensions than the more complex shape space tested here, and our findings may therefore reveal important boundary conditions of feature distribution learning.

You see what you look for: Targets and distractors in visual search can cause opposing serial dependencies.

Visual perception is, at any given moment, strongly influenced by its temporal context-what stimuli have recently been perceived and in what surroundings. We have previously shown that to-be-ignored items produce a bias upon subsequent perceptual decisions that acts in parallel with other biases induced by attended items. However, our previous investigations were confined to biases upon the perceived orientation of a visual search target, and it is unclear whether these biases influence perceptual decisions in a more general sense. Here, we test whether the biases from visual search targets and distractors affect the perceived orientation of a neutral test line, one that is neither a target nor a distractor. To do so, we asked participants to search for an oddly oriented line among distractors and report its location for a few trials and next presented a test line irrelevant to the search task. Participants were asked to report the orientation of the test line. Our results indicate that in tasks involving visual search, targets induce a positive bias upon a neutral test line if their orientations are similar, whereas distractors produce an attractive bias for similar test lines and a repulsive bias if the orientations of the test line and the average orientation of the distractors are far apart in feature space. In sum, our results show that both attentional role and proximity in feature space between previous and current stimuli determine the direction of biases in perceptual decisions.

What kind of empirical evidence is needed for probabilistic mental representations? An example from visual perception.

Recent accounts of perception and cognition propose that the brain represents information probabilistically. While this assumption is common, empirical support for such probabilistic representations in perception has recently been criticized. Here, we evaluate these criticisms and present an account based on a recently developed psychophysical methodology, Feature Distribution Learning (FDL), which provides promising evidence for probabilistic representations by avoiding these criticisms. The method uses priming and role-reversal effects in visual search. Observers' search times reveal the structure of perceptual representations, in which the probability distribution of distractor features is encoded. We explain how FDL results provide evidence for a stronger notion of representation that relies on structural correspondence between stimulus uncertainty and perceptual representations, rather than a mere co-variation between the two. Moreover, such an account allows us to demonstrate what kind of empirical evidence is needed to support probabilistic representations as posited in current probabilistic Bayesian theories of perception.

Testing temporal integration of feature probability distributions using role-reversal effects in visual search.

The visual system is sensitive to statistical properties of complex scenes and can encode feature probability distributions in detail. But does the brain use these statistics to build probabilistic models of the ever-changing visual input? To investigate this, we examined how observers temporally integrate two different orientation distributions from sequentially presented visual search trials. If the encoded probabilistic information is used in a Bayesian optimal way, observers should weigh more reliable information more strongly, such as feature distributions with low variance. We therefore manipulated the variance of the two feature distributions. Participants performed sequential odd-one-out visual search for an oddly oriented line among distractors. During successive learning trials, the distractor orientations were sampled from two different Gaussian distributions on alternating trials. Then, observers performed a 'test trial' where the orientations of the target and distractors were switched, allowing us to assess observer's internal representation of distractor distributions based on changes in response times. In three experiments we observed that observer's search times on test trials depended mainly on the very last learning trial, indicating a strong recency effect. Since temporal integration has been previously observed with this method, we conclude that when the input is unreliable, the visual system relies more on the most recent stimulus. This indicates that the visual system prefers to utilize sensory history when the statistical properties of the environment are relatively stable.

Dual strategies in human confidence judgments.

Although confidence is commonly believed to be an essential element in decision-making, it remains unclear what gives rise to one's sense of confidence. Recent Bayesian theories propose that confidence is computed, in part, from the degree of uncertainty in sensory evidence. Alternatively, observers can use physical properties of the stimulus as a heuristic to confidence. In the current study, we developed ideal observer models for either hypothesis and compared their predictions against human data obtained from psychophysical experiments. Participants reported the orientation of a stimulus, and their confidence in this estimate, under varying levels of internal and external noise. As predicted by the Bayesian model, we found a consistent link between confidence and behavioral variability for a given stimulus orientation. Confidence was higher when orientation estimates were more precise, for both internal and external sources of noise. However, we observed the inverse pattern when comparing between stimulus orientations: although observers gave more precise orientation estimates for cardinal orientations (a phenomenon known as the oblique effect), they were more confident about oblique orientations. We show that these results are well explained by a strategy to confidence that is based on the perceived amount of noise in the stimulus. Altogether, our results suggest that confidence is not always computed from the degree of uncertainty in one's perceptual evidence but can instead be based on visual cues that function as simple Heuristics to confidence.

Structural and contextual priors affect visual search in children with and without autism.

Bayesian predictive coding theories of autism spectrum disorder propose that impaired acquisition or a broader shape of prior probability distributions lies at the core of the condition. However, we still know very little about how probability distributions are learned and encoded by children, let alone children with autism. Here, we take advantage of a recently developed distribution learning paradigm to characterize how children with and without autism acquire information about probability distributions. Twenty-four autistic and 25-matched neurotypical children searched for an odd-one-out target among a set of distractor lines with orientations sampled from a Gaussian distribution repeated across multiple trials to allow for learning of the parameters (mean and variance) of the distribution. We could measure the width (variance) of the participant's encoded distribution by introducing a target-distractor role-reversal while varying the similarity between target and previous distractor mean. Both groups performed similarly on the visual search task and learned the distractor distribution to a similar extent. However, the variance learned was much broader than the one presented, consistent with less informative priors in children irrespective of autism diagnosis. These findings have important implications for Bayesian accounts of perception throughout development, and Bayesian accounts of autism specifically. LAY SUMMARY: Recent theories about the underlying cognitive mechanisms of autism propose that the way autistic individuals estimate variability or uncertainty in their perceptual environment may differ from how typical individuals do so. Children had to search an oddly tilted line in a set of lines pointing in different directions, and based on their response times we examined how they learned about the variability in a set of objects. We found that autistic children learn variability as well as typical children, but both groups learn with less precision than typical adults do on the same task.

Dissociating implicit and explicit ensemble representations reveals the limits of visual perception and the richness of behavior.

Our senses provide us with a rich experience of a detailed visual world, yet the empirical results seem to suggest severe limitations on our ability to perceive and remember. In recent attempts to reconcile the contradiction between what is experienced and what can be reported, it has been argued that the visual world is condensed to a set of summary statistics, explaining both the rich experience and the sparse reports. Here, we show that explicit reports of summary statistics underestimate the richness of ensemble perception. Our observers searched for an odd-one-out target among heterogeneous distractors and their representation of distractor characteristics was tested explicitly or implicitly. Observers could explicitly distinguish distractor sets with different mean and variance, but not differently-shaped probability distributions. In contrast, the implicit assessment revealed that the visual system encodes the mean, the variance, and even the shape of feature distributions. Furthermore, explicit measures had common noise sources that distinguished them from implicit measures. This suggests that explicit judgments of stimulus ensembles underestimate the richness of visual representations. We conclude that feature distributions are encoded in rich detail and can guide behavior implicitly, even when the information available for explicit summary judgments is coarse and limited.

Optimizing perception: Attended and ignored stimuli create opposing perceptual biases.

Humans have remarkable abilities to construct a stable visual world from continuously changing input. There is increasing evidence that momentary visual input blends with previous input to preserve perceptual continuity. Most studies have shown that such influences can be traced to characteristics of the attended object at a given moment. Little is known about the role of ignored stimuli in creating this continuity. This is important since while some input is selected for processing, other input must be actively ignored for efficient selection of the task-relevant stimuli. We asked whether attended targets and actively ignored distractor stimuli in an odd-one-out search task would bias observers' perception differently. Our observers searched for an oddly oriented line among distractors and were occasionally asked to report the orientation of the last visual search target they saw in an adjustment task. Our results show that at least two opposite biases from past stimuli influence current perception: A positive bias caused by serial dependence pulls perception of the target toward the previous target features, while a negative bias induced by the to-be-ignored distractor features pushes perception of the target away from the distractor distribution. Our results suggest that to-be-ignored items produce a perceptual bias that acts in parallel with other biases induced by attended items to optimize perception. Our results are the first to demonstrate how actively ignored information facilitates continuity in visual perception.

Encoding perceptual ensembles during visual search in peripheral vision.

Observers can learn complex statistical properties of visual ensembles, such as their probability distributions. Even though ensemble encoding is considered critical for peripheral vision, whether observers learn such distributions in the periphery has not been studied. Here, we used a visual search task to investigate how the shape of distractor distributions influences search performance and ensemble encoding in peripheral and central vision. Observers looked for an oddly oriented bar among distractors taken from either uniform or Gaussian orientation distributions with the same mean and range. The search arrays were either presented in the foveal or peripheral visual fields. The repetition and role reversal effects on search times revealed observers' internal model of distractor distributions. Our results showed that the shape of the distractor distribution influenced search times only in foveal, but not in peripheral search. However, role reversal effects revealed that the shape of the distractor distribution could be encoded peripherally depending on the interitem spacing in the search array. Our results suggest that, although peripheral vision might rely heavily on summary statistical representations of feature distributions, it can also encode information about the distributions themselves.

No Advantage for Separating Overt and Covert Attention in Visual Search.

We move our eyes roughly three times every second while searching complex scenes, but covert attention helps to guide where we allocate those overt fixations. Covert attention may be allocated reflexively or voluntarily, and speeds the rate of information processing at the attended location. Reducing access to covert attention hinders performance, but it is not known to what degree the locus of covert attention is tied to the current gaze position. We compared visual search performance in a traditional gaze-contingent display, with a second task where a similarly sized contingent window is controlled with a mouse, allowing a covert aperture to be controlled independently by overt gaze. Larger apertures improved performance for both the mouse- and gaze-contingent trials, suggesting that covert attention was beneficial regardless of control type. We also found evidence that participants used the mouse-controlled aperture somewhat independently of gaze position, suggesting that participants attempted to untether their covert and overt attention when possible. This untethering manipulation, however, resulted in an overall cost to search performance, a result at odds with previous results in a change blindness paradigm. Untethering covert and overt attention may therefore have costs or benefits depending on the task demands in each case.

The Confidence Database.

Understanding how people rate their confidence is critical for the characterization of a wide range of perceptual, memory, motor and cognitive processes. To enable the continued exploration of these processes, we created a large database of confidence studies spanning a broad set of paradigms, participant populations and fields of study. The data from each study are structured in a common, easy-to-use format that can be easily imported and analysed using multiple software packages. Each dataset is accompanied by an explanation regarding the nature of the collected data. At the time of publication, the Confidence Database (which is available at https://osf.io/s46pr/) contained 145 datasets with data from more than 8,700 participants and almost 4 million trials. The database will remain open for new submissions indefinitely and is expected to continue to grow. Here we show the usefulness of this large collection of datasets in four different analyses that provide precise estimations of several foundational confidence-related effects.

Dynamics of visual attention revealed in foraging tasks.

Visual search tasks play a key role in theories of visual attention. But single-target search tasks may provide only a snapshot of attentional orienting. Foraging tasks with multiple targets of different types arguably provide a closer analogy to everyday attentional processing. Set-size effects have in the literature formed the basis for inferring how attention operates during visual search. We therefore measured the effects of absolute set-size (constant target-distractor ratio) and relative set-size (constant set-size but target-distractor ratio varies) on foraging patterns during "feature" foraging (targets differed from distractors on a single feature) and "conjunction" foraging (targets differed from distractors on a combination of two features). Patterns of runs of same target-type selection were similar regardless of whether absolute or relative set-size varied: long sequential runs during conjunction foraging but rapid switching between target types during feature foraging. But although foraging strategies differed between feature and conjunction foraging, surprisingly, intertarget times throughout foraging trials did not differ much between the conditions. Typical response time by set-size patterns for single-target visual search tasks were only observed for the last target during foraging. Furthermore, the foraging patterns within trials involved several distinct phases, that may serve as markers of particular attentional operations. Foraging tasks provide a remarkably intricate picture of attentional selection, far more detailed than traditional single-target visual search tasks, and well-known theories of visual attention have difficulty accounting for key aspects of the observed foraging patterns. Finally, we discuss how theoretical conceptions of attention could be modified to account for these effects.

Probabilistic rejection templates in visual working memory.

Our interactions with the visual world are guided by attention and visual working memory. Things that we look for and those we ignore are stored as templates that reflect our goals and the tasks at hand. The nature of such templates has been widely debated. A recent proposal is that these templates can be thought of as probabilistic representations of task-relevant features. Crucially, such probabilistic templates should accurately reflect feature probabilities in the environment. Here we ask whether observers can quickly form a correct internal model of a complex (bimodal) distribution of distractor features. We assessed observers' representations by measuring the slowing of visual search when target features unexpectedly match a distractor template. Distractor stimuli were heterogeneous, randomly drawn on each trial from a bimodal probability distribution. Using two targets on each trial, we tested whether observers encode the full distribution, only one peak of it, or the average of the two peaks. Search was slower when the two targets corresponded to the two modes of a previous distractor distribution than when one target was at one of the modes and another between them or outside the distribution range. Furthermore, targets on the modes were reported later than targets between the modes that, in turn, were reported later than targets outside this range. This shows that observers use a correct internal model, representing both distribution modes using templates based on the full probability distribution rather than just one peak or simple summary statistics. The findings further confirm that performance in odd-one out search with repeated distractors cannot be described by a simple decision rule. Our findings indicate that probabilistic visual working memory templates guiding attention, dynamically adapt to task requirements, accurately reflecting the probabilistic nature of the input.

Representing color and orientation ensembles: Can observers learn multiple feature distributions?

Objects have a variety of different features that can be represented as probability distributions. Recent findings show that in addition to mean and variance, the visual system can also encode the shape of feature distributions for features like color or orientation. In an odd-one-out search task we investigated observers' ability to encode two feature distributions simultaneously. Our stimuli were defined by two distinct features (color and orientation) while only one was relevant to the search task. We investigated whether the irrelevant feature distribution influences learning of the task-relevant distribution and whether observers also encode the irrelevant distribution. Although considerable learning of feature distributions occurred, especially for color, our results also suggest that adding a second irrelevant feature distribution negatively affected the encoding of the relevant one and that little learning of the irrelevant distribution occurred. There was also an asymmetry between the two different features: Searching for the oddly oriented target was more difficult than searching for the oddly colored target, which was reflected in worse learning of the color distribution. Overall, the results demonstrate that it is possible to encode information about two feature distributions simultaneously but also reveal considerable limits to this encoding.

Expectations and perceptual priming in a visual search task: Evidence from eye movements and behavior.

An extensive amount of research indicates that repeating target and distractor features facilitates pop-out search while switching these features slows the search. Following the seminal study by Maljkovic and Nakayama (1994), this "priming of pop-out" effect (PoP) has been widely described as an automatic bottom-up process that is independent of the observers' expectations. At the same time, numerous studies highlight the crucial role of expectations in visual attention deployment. Our experiment shows that in contrast to previous claims, PoP in a classic color singleton search task is a mix of automatic processing and expectations. Participants searched for a uniquely colored diamond among 2 same-colored distractors. Target color sequences were either predictable (e.g., 2 red-target-green-distractors trials, followed by 2 green-target-red-distractors trials, and so on) or random. Responses were faster in predictable color sequences than randomly changing ones with equal number of repetitions of target colors on preceding trials. Analyses of observers' eye movements showed that predictability of target color affected both latency and accuracy of the first saccade during a search trial. Our results support the idea that PoP is governed not only by automatic effects from previous target or distractor features but also by top-down expectations. (PsycINFO Database Record (c) 2019 APA, all rights reserved).

A different kind of pain: affective valence of errors and incongruence.

People hiss and swear when they make errors, frown and swear again when they encounter conflicting information. Such error- and conflict-related signs of negative affect are found even when there is no time pressure or external reward and the task itself is very simple. Previous studies, however, provide inconsistent evidence regarding the affective consequences of resolved conflicts, that is, conflicts that resulted in correct responses. We tested whether response accuracy in the Eriksen flanker task will moderate the effect of trial incongruence using affective priming to measure positive and negative affect. We found that responses to incongruent trials elicit positive affect irrespective of their accuracy. Errors, in turn, result in negative affect irrespective of trial congruence. The effects of conflicts and errors do not interact and affect different dimensions of affective priming. Conflicts change the speed of evaluative categorisation while errors are reflected in categorisation accuracy. We discuss the findings in light of the "reward value and prediction" model and the "affect as a feedback for predictions" framework and consider the possible mechanisms behind the divergent effects.