Expertise increases planning depth in human gameplay.

A hallmark of human intelligence is the ability to plan multiple steps into the future1,2. Despite decades of research3-5, it is still debated whether skilled decision-makers plan more steps ahead than novices6-8. Traditionally, the study of expertise in planning has used board games such as chess, but the complexity of these games poses a barrier to quantitative estimates of planning depth. Conversely, common planning tasks in cognitive science often have a lower complexity9,10 and impose a ceiling for the depth to which any player can plan. Here we investigate expertise in a complex board game that offers ample opportunity for skilled players to plan deeply. We use model fitting methods to show that human behaviour can be captured using a computational cognitive model based on heuristic search. To validate this model, we predict human choices, response times and eye movements. We also perform a Turing test and a reconstruction experiment. Using the model, we find robust evidence for increased planning depth with expertise in both laboratory and large-scale mobile data. Experts memorize and reconstruct board features more accurately. Using complex tasks combined with precise behavioural modelling might expand our understanding of human planning and help to bridge the gap with progress in artificial intelligence.

Normalization by orientation-tuned surround in human V1-V3.

An influential account of neuronal responses in primary visual cortex is the normalized energy model. This model is often implemented as a multi-stage computation. The first stage is linear filtering. The second stage is the extraction of contrast energy, whereby a complex cell computes the squared and summed outputs of a pair of the linear filters in quadrature phase. The third stage is normalization, in which a local population of complex cells mutually inhibit one another. Because the population includes cells tuned to a range of orientations and spatial frequencies, the result is that the responses are effectively normalized by the local stimulus contrast. Here, using evidence from human functional MRI, we show that the classical model fails to account for the relative responses to two classes of stimuli: straight, parallel, band-passed contours (gratings), and curved, band-passed contours (snakes). The snakes elicit fMRI responses that are about twice as large as the gratings, yet a traditional divisive normalization model predicts responses that are about the same. Motivated by these observations and others from the literature, we implement a divisive normalization model in which cells matched in orientation tuning ("tuned normalization") preferentially inhibit each other. We first show that this model accounts for differential responses to these two classes of stimuli. We then show that the model successfully generalizes to other band-pass textures, both in V1 and in extrastriate cortex (V2 and V3). We conclude that even in primary visual cortex, complex features of images such as the degree of heterogeneity, can have large effects on neural responses.

Humans incorporate trial-to-trial working memory uncertainty into rewarded decisions.

Working memory (WM) plays an important role in action planning and decision making; however, both the informational content of memory and how that information is used in decisions remain poorly understood. To investigate this, we used a color WM task in which subjects viewed colored stimuli and reported both an estimate of a stimulus color and a measure of memory uncertainty, obtained through a rewarded decision. Reported memory uncertainty is correlated with memory error, showing that people incorporate their trial-to-trial memory quality into rewarded decisions. Moreover, memory uncertainty can be combined with other sources of information; after inducing expectations (prior beliefs) about stimuli probabilities, we found that estimates became shifted toward expected colors, with the shift increasing with reported uncertainty. The data are best fit by models in which people incorporate their trial-to-trial memory uncertainty with potential rewards and prior beliefs. Our results suggest that WM represents uncertainty information, and that this can be combined with prior beliefs. This highlights the potential complexity of WM representations and shows that rewarded decision can be a powerful tool for examining WM and informing and constraining theoretical, computational, and neurobiological models of memory.

Neural coding of uncertainty and probability.

Organisms must act in the face of sensory, motor, and reward uncertainty stemming from a pandemonium of stochasticity and missing information. In many tasks, organisms can make better decisions if they have at their disposal a representation of the uncertainty associated with task-relevant variables. We formalize this problem using Bayesian decision theory and review recent behavioral and neural evidence that the brain may use knowledge of uncertainty, confidence, and probability.

An uncertainty-based model of the effects of fixation on choice.

When people view a consumable item for a longer amount of time, they choose it more frequently; this also seems to be the direction of causality. The leading model of this effect is a drift-diffusion model with a fixation-based attentional bias. Here, we propose an explicitly Bayesian account for the same data. This account is based on the notion that the brain builds a posterior belief over the value of an item in the same way it would over a sensory variable. As the agent gathers evidence about the item from sensory observations and from retrieved memories, the posterior distribution narrows. We further postulate that the utility of an item is a weighted sum of the posterior mean and the negative posterior standard deviation, with the latter accounting for risk aversion. Fixating for longer can increase or decrease the posterior mean, but will inevitably lower the posterior standard deviation. This model fits the data better than the original attentional drift-diffusion model but worse than a variant with a collapsing bound. We discuss the often overlooked technical challenges in fitting models simultaneously to choice and response time data in the absence of an analytical expression. Our results hopefully contribute to emerging accounts of valuation as an inference process.

Point-estimating observer models for latent cause detection.

The spatial distribution of visual items allows us to infer the presence of latent causes in the world. For instance, a spatial cluster of ants allows us to infer the presence of a common food source. However, optimal inference requires the integration of a computationally intractable number of world states in real world situations. For example, optimal inference about whether a common cause exists based on N spatially distributed visual items requires marginalizing over both the location of the latent cause and 2N possible affiliation patterns (where each item may be affiliated or non-affiliated with the latent cause). How might the brain approximate this inference? We show that subject behaviour deviates qualitatively from Bayes-optimal, in particular showing an unexpected positive effect of N (the number of visual items) on the false-alarm rate. We propose several "point-estimating" observer models that fit subject behaviour better than the Bayesian model. They each avoid a costly computational marginalization over at least one of the variables of the generative model by "committing" to a point estimate of at least one of the two generative model variables. These findings suggest that the brain may implement partially committal variants of Bayesian models when detecting latent causes based on complex real world data.

Unbiased and efficient log-likelihood estimation with inverse binomial sampling.

The fate of scientific hypotheses often relies on the ability of a computational model to explain the data, quantified in modern statistical approaches by the likelihood function. The log-likelihood is the key element for parameter estimation and model evaluation. However, the log-likelihood of complex models in fields such as computational biology and neuroscience is often intractable to compute analytically or numerically. In those cases, researchers can often only estimate the log-likelihood by comparing observed data with synthetic observations generated by model simulations. Standard techniques to approximate the likelihood via simulation either use summary statistics of the data or are at risk of producing substantial biases in the estimate. Here, we explore another method, inverse binomial sampling (IBS), which can estimate the log-likelihood of an entire data set efficiently and without bias. For each observation, IBS draws samples from the simulator model until one matches the observation. The log-likelihood estimate is then a function of the number of samples drawn. The variance of this estimator is uniformly bounded, achieves the minimum variance for an unbiased estimator, and we can compute calibrated estimates of the variance. We provide theoretical arguments in favor of IBS and an empirical assessment of the method for maximum-likelihood estimation with simulation-based models. As case studies, we take three model-fitting problems of increasing complexity from computational and cognitive neuroscience. In all problems, IBS generally produces lower error in the estimated parameters and maximum log-likelihood values than alternative sampling methods with the same average number of samples. Our results demonstrate the potential of IBS as a practical, robust, and easy to implement method for log-likelihood evaluation when exact techniques are not available.

The role of sensory uncertainty in simple contour integration.

Perceptual organization is the process of grouping scene elements into whole entities. A classic example is contour integration, in which separate line segments are perceived as continuous contours. Uncertainty in such grouping arises from scene ambiguity and sensory noise. Some classic Gestalt principles of contour integration, and more broadly, of perceptual organization, have been re-framed in terms of Bayesian inference, whereby the observer computes the probability that the whole entity is present. Previous studies that proposed a Bayesian interpretation of perceptual organization, however, have ignored sensory uncertainty, despite the fact that accounting for the current level of perceptual uncertainty is one of the main signatures of Bayesian decision making. Crucially, trial-by-trial manipulation of sensory uncertainty is a key test to whether humans perform near-optimal Bayesian inference in contour integration, as opposed to using some manifestly non-Bayesian heuristic. We distinguish between these hypotheses in a simplified form of contour integration, namely judging whether two line segments separated by an occluder are collinear. We manipulate sensory uncertainty by varying retinal eccentricity. A Bayes-optimal observer would take the level of sensory uncertainty into account-in a very specific way-in deciding whether a measured offset between the line segments is due to non-collinearity or to sensory noise. We find that people deviate slightly but systematically from Bayesian optimality, while still performing "probabilistic computation" in the sense that they take into account sensory uncertainty via a heuristic rule. Our work contributes to an understanding of the role of sensory uncertainty in higher-order perception.

Humans incorporate attention-dependent uncertainty into perceptual decisions and confidence.

Perceptual decisions are better when they take uncertainty into account. Uncertainty arises not only from the properties of sensory input but also from cognitive sources, such as different levels of attention. However, it is unknown whether humans appropriately adjust for such cognitive sources of uncertainty during perceptual decision-making. Here we show that, in a task in which uncertainty is relevant for performance, human categorization and confidence decisions take into account uncertainty related to attention. We manipulated uncertainty in an orientation categorization task from trial to trial using only an attentional cue. The categorization task was designed to disambiguate decision rules that did or did not depend on attention. Using formal model comparison to evaluate decision behavior, we found that category and confidence decision boundaries shifted as a function of attention in an approximately Bayesian fashion. This means that the observer's attentional state on each trial contributed probabilistically to the decision computation. This responsiveness of an observer's decisions to attention-dependent uncertainty should improve perceptual decisions in natural vision, in which attention is unevenly distributed across a scene.

Uncertainty is maintained and used in working memory.

What are the contents of working memory? In both behavioral and neural computational models, a working memory representation is typically described by a single number, namely, a point estimate of a stimulus. Here, we asked if people also maintain the uncertainty associated with a memory and if people use this uncertainty in subsequent decisions. We collected data in a two-condition orientation change detection task; while both conditions measured whether people used memory uncertainty, only one required maintaining it. For each condition, we compared an optimal Bayesian observer model, in which the observer uses an accurate representation of uncertainty in their decision, to one in which the observer does not. We find that this "Use Uncertainty" model fits better for all participants in both conditions. In the first condition, this result suggests that people use uncertainty optimally in a working memory task when that uncertainty information is available at the time of decision, confirming earlier results. Critically, the results of the second condition suggest that this uncertainty information was maintained in working memory. We test model variants and find that our conclusions do not depend on our assumptions about the observer's encoding process, inference process, or decision rule. Our results provide evidence that people have uncertainty that reflects their memory precision on an item-specific level, maintain this information over a working memory delay, and use it implicitly in a way consistent with an optimal observer. These results challenge existing computational models of working memory to update their frameworks to represent uncertainty.

Human online adaptation to changes in prior probability.

Optimal sensory decision-making requires the combination of uncertain sensory signals with prior expectations. The effect of prior probability is often described as a shift in the decision criterion. Can observers track sudden changes in probability? To answer this question, we used a change-point detection paradigm that is frequently used to examine behavior in changing environments. In a pair of orientation-categorization tasks, we investigated the effects of changing probabilities on decision-making. In both tasks, category probability was updated using a sample-and-hold procedure: probability was held constant for a period of time before jumping to another probability state that was randomly selected from a predetermined set of probability states. We developed an ideal Bayesian change-point detection model in which the observer marginalizes over both the current run length (i.e., time since last change) and the current category probability. We compared this model to various alternative models that correspond to different strategies-from approximately Bayesian to simple heuristics-that the observers may have adopted to update their beliefs about probabilities. While a number of models provided decent fits to the data, model comparison favored a model in which probability is estimated following an exponential averaging model with a bias towards equal priors, consistent with a conservative bias, and a flexible variant of the Bayesian change-point detection model with incorrect beliefs. We interpret the former as a simpler, more biologically plausible explanation suggesting that the mechanism underlying change of decision criterion is a combination of on-line estimation of prior probability and a stable, long-term equal-probability prior, thus operating at two very different timescales.

Explaining the effects of distractor statistics in visual search.

Visual search, the task of detecting or locating target items among distractor items in a visual scene, is an important function for animals and humans. Different theoretical accounts make differing predictions for the effects of distractor statistics. Here we use a task in which we parametrically vary distractor items, allowing for a simultaneously fine-grained and comprehensive study of distractor statistics. We found effects of target-distractor similarity, distractor variability, and an interaction between the two, although the effect of the interaction on performance differed from the one expected. To explain these findings, we constructed computational process models that make trial-by-trial predictions for behavior based on the stimulus presented. These models, including a Bayesian observer model, provided excellent accounts of both the qualitative and quantitative effects of distractor statistics, as well as of the effect of changing the statistics of the environment (in the form of distractors being drawn from a different distribution). We conclude with a broader discussion of the role of computational process models in the understanding of visual search.

Multiple conceptions of resource rationality.

Resource rationality holds great promise as a unifying principle across theories in neuroscience, cognitive science, and economics. The target article clearly lays out this potential for unification. However, resource-rational models are more diverse and less easily unified than might appear from the target article. Here, we explore some of that diversity.

Comparing Bayesian and non-Bayesian accounts of human confidence reports.

Humans can meaningfully report their confidence in a perceptual or cognitive decision. It is widely believed that these reports reflect the Bayesian probability that the decision is correct, but this hypothesis has not been rigorously tested against non-Bayesian alternatives. We use two perceptual categorization tasks in which Bayesian confidence reporting requires subjects to take sensory uncertainty into account in a specific way. We find that subjects do take sensory uncertainty into account when reporting confidence, suggesting that brain areas involved in reporting confidence can access low-level representations of sensory uncertainty, a prerequisite of Bayesian inference. However, behavior is not fully consistent with the Bayesian hypothesis and is better described by simple heuristic models that use uncertainty in a non-Bayesian way. Both conclusions are robust to changes in the uncertainty manipulation, task, response modality, model comparison metric, and additional flexibility in the Bayesian model. Our results suggest that adhering to a rational account of confidence behavior may require incorporating implementational constraints.

Bayesian comparison of explicit and implicit causal inference strategies in multisensory heading perception.

The precision of multisensory perception improves when cues arising from the same cause are integrated, such as visual and vestibular heading cues for an observer moving through a stationary environment. In order to determine how the cues should be processed, the brain must infer the causal relationship underlying the multisensory cues. In heading perception, however, it is unclear whether observers follow the Bayesian strategy, a simpler non-Bayesian heuristic, or even perform causal inference at all. We developed an efficient and robust computational framework to perform Bayesian model comparison of causal inference strategies, which incorporates a number of alternative assumptions about the observers. With this framework, we investigated whether human observers' performance in an explicit cause attribution and an implicit heading discrimination task can be modeled as a causal inference process. In the explicit causal inference task, all subjects accounted for cue disparity when reporting judgments of common cause, although not necessarily all in a Bayesian fashion. By contrast, but in agreement with previous findings, data from the heading discrimination task only could not rule out that several of the same observers were adopting a forced-fusion strategy, whereby cues are integrated regardless of disparity. Only when we combined evidence from both tasks we were able to rule out forced-fusion in the heading discrimination task. Crucially, findings were robust across a number of variants of models and analyses. Our results demonstrate that our proposed computational framework allows researchers to ask complex questions within a rigorous Bayesian framework that accounts for parameter and model uncertainty.

Limitations of Proposed Signatures of Bayesian Confidence.

The Bayesian model of confidence posits that confidence reflects the observer's posterior probability that the decision is correct. Hangya, Sanders, and Kepecs (2016) have proposed that researchers can test the Bayesian model by deriving qualitative signatures of Bayesian confidence (i.e., patterns that one would expect to see if an observer were Bayesian) and looking for those signatures in human or animal data. We examine two proposed signatures, showing that their derivations contain hidden assumptions that limit their applicability and that they are neither necessary nor sufficient conditions for Bayesian confidence. One signature is an average confidence of 0.75 on trials with neutral evidence. This signature holds only when class-conditioned stimulus distributions do not overlap and when internal noise is very low. Another signature is that as stimulus magnitude increases, confidence increases on correct trials but decreases on incorrect trials. This divergence signature holds only when stimulus distributions do not overlap or when noise is high. Navajas et al. (2017) have proposed an alternative form of this signature; we find no indication that this alternative form is expected under Bayesian confidence. Our observations give us pause about the usefulness of the qualitative signatures of Bayesian confidence. To determine the nature of the computations underlying confidence reports, there may be no shortcut to quantitative model comparison.

Identifying suboptimalities with factorial model comparison.

Given the many types of suboptimality in perception, I ask how one should test for multiple forms of suboptimality at the same time - or, more generally, how one should compare process models that can differ in any or all of the multiple components. In analogy to factorial experimental design, I advocate for factorial model comparison.

Visual short-term memory for oriented, colored objects.

A central question in the study of visual short-term memory (VSTM) has been whether its basic units are objects or features. Most studies addressing this question have used change detection tasks in which the feature value before the change is highly discriminable from the feature value after the change. This approach assumes that memory noise is negligible, which recent work has shown not to be the case. Here, we investigate VSTM for orientation and color within a noisy-memory framework, using change localization with a variable magnitude of change. A specific consequence of the noise is that it is necessary to model the inference (decision) stage. We find that (a) orientation and color have independent pools of memory resource (consistent with classic results); (b) an irrelevant feature dimension is either encoded but ignored during decision-making, or encoded with low precision and taken into account during decision-making; and (c) total resource available in a given feature dimension is lower in the presence of task-relevant stimuli that are neutral in that feature dimension. We propose a framework in which feature resource comes both in packaged and in targeted form.

The effects of delay duration on visual working memory for orientation.

We used a delayed-estimation paradigm to characterize the joint effects of set size (one, two, four, or six) and delay duration (1, 2, 3, or 6 s) on visual working memory for orientation. We conducted two experiments: one with delay durations blocked, another with delay durations interleaved. As dependent variables, we examined four model-free metrics of dispersion as well as precision estimates in four simple models. We tested for effects of delay time using analyses of variance, linear regressions, and nested model comparisons. We found significant effects of set size and delay duration on both model-free and model-based measures of dispersion. However, the effect of delay duration was much weaker than that of set size, dependent on the analysis method, and apparent in only a minority of subjects. The highest forgetting slope found in either experiment at any set size was a modest 1.14°/s. As secondary results, we found a low rate of nontarget reports, and significant estimation biases towards oblique orientations (but no dependence of their magnitude on either set size or delay duration). Relative stability of working memory even at higher set sizes is consistent with earlier results for motion direction and spatial frequency. We compare with a recent study that performed a very similar experiment.

Orientation-dependent biases in length judgments of isolated stimuli.

Vertical line segments tend to be perceived as longer than horizontal ones of the same length, but this may in part be due to configuration effects. To minimize such effects, we used isolated line segments in a two-interval, forced choice paradigm, not limiting ourselves to horizontal and vertical. We fitted psychometric curves using a Bayesian method that assumes that, for a given subject, the lapse rate is the same across all conditions. The closer a line segment's orientation was to vertical, the longer it was perceived to be. Moreover, subjects tended to report the standard line (in the second interval) as longer. The data were well described by a model that contains both an orientation-dependent and an interval-dependent multiplicative bias. Using this model, we estimated that a vertical line was on average perceived as 9.2% ± 2.1% longer than a horizontal line, and a second-interval line was on average perceived as 2.4% ± 0.9% longer than a first-interval line. Moving from a descriptive to an explanatory model, we hypothesized that anisotropy in the polar angle of lines in three dimensions underlies the horizontal-vertical illusion, specifically, that line segments more often have a polar angle of 90° (corresponding to the ground plane) than any other polar angle. This model qualitatively accounts not only for the empirical relationship between projected length and projected orientation that predicts the horizontal-vertical illusion, but also for the empirical distribution of projected orientation in photographs of natural scenes and for paradoxical results reported earlier for slanted surfaces.