Confirmation Bias through Selective Use of Evidence in Human Cortex.

Decision-makers often process new evidence selectively, depending on their current beliefs about the world. We asked whether such confirmation biases result from biases in the encoding of sensory evidence in the brain, or alternatively in the utilization of encoded evidence for behavior. Human participants estimated the source of a sequence of visual-spatial evidence samples while we measured cortical population activity with magnetoencephalography (MEG). Halfway through the sequence, participants were prompted to judge the more likely source category. Their processing of subsequent evidence depended on its consistency with the previously chosen category, but the encoding of evidence in cortical activity did not. Instead, the encoded evidence in parietal and primary visual cortex contributed less to the estimation report when that evidence was inconsistent with the previous choice. We conclude that confirmation bias originates from the way in which decision-makers utilize information encoded in the brain. This provides room for deliberative control.

Sensory perception is a holistic inference process.

Sensory perception is widely considered an inference process that reflects the best guess of a stimulus feature based on uncertain sensory information. Here we challenge this reductionist view and propose that perception is rather a holistic inference process that operates not only at the feature but jointly across all levels of the representational hierarchy. We test this hypothesis in the context of a commonly used psychophysical matching task in which subjects are asked to report their perceived orientation of a test stimulus by adjusting a probe stimulus (method-of-adjustment). We introduce a holistic matching model that assumes that subjects' reports reflect an optimal match between the test and probe stimulus, both in terms of their inferred feature (orientation) and also their higher level representation (orientation category). Validation against several existing data sets demonstrates that the model accurately and comprehensively predicts subjects' response behavior and outperforms previous models both qualitatively and quantitatively. Moreover, the model generalizes to other feature domains and offers an alternative account for categorical color perception. Our results suggest that categorical effects in sensory perception are ubiquitous and can be parsimoniously explained as optimal behavior based on holistic sensory representations. (PsycInfo Database Record (c) 2024 APA, all rights reserved).

Efficient sensory encoding predicts robust averaging.

Not every item in a stimulus ensemble equally contributes to the perceived ensemble average. Rather, items with feature values close to the ensemble mean (inlying items) contribute stronger compared to those items whose feature values are further away from the mean (outlying items). This nonuniform weighting process, named robust averaging, has been interpreted as evidence against an optimal integration of sensory information. Here, however, we show that robust averaging naturally emerges from an optimal integration process when sensory encoding is efficiently adapted to the ensemble statistics in the experiment. We demonstrate that such a model can accurately fit several existing datasets showing robust perceptual averaging in discriminating low-level stimulus features such as orientation. Across various feature domains, our model accurately predicts subjects' decision accuracy and nonuniform weighting profile, and both their dependency on the specific stimulus distribution in the experiments. Our results suggest that the human visual system forms efficient sensory representations on short time-scales to improve overall decision performance.

Efficient neural codes naturally emerge through gradient descent learning.

Human sensory systems are more sensitive to common features in the environment than uncommon features. For example, small deviations from the more frequently encountered horizontal orientations can be more easily detected than small deviations from the less frequent diagonal ones. Here we find that artificial neural networks trained to recognize objects also have patterns of sensitivity that match the statistics of features in images. To interpret these findings, we show mathematically that learning with gradient descent in neural networks preferentially creates representations that are more sensitive to common features, a hallmark of efficient coding. This effect occurs in systems with otherwise unconstrained coding resources, and additionally when learning towards both supervised and unsupervised objectives. This result demonstrates that efficient codes can naturally emerge from gradient-like learning.

Prior Expectations in Visual Speed Perception Predict Encoding Characteristics of Neurons in Area MT.

Bayesian inference provides an elegant theoretical framework for understanding the characteristic biases and discrimination thresholds in visual speed perception. However, the framework is difficult to validate because of its flexibility and the fact that suitable constraints on the structure of the sensory uncertainty have been missing. Here, we demonstrate that a Bayesian observer model constrained by efficient coding not only well explains human visual speed perception but also provides an accurate quantitative account of the tuning characteristics of neurons known for representing visual speed. Specifically, we found that the population coding accuracy for visual speed in area MT ("neural prior") is precisely predicted by the power-law, slow-speed prior extracted from fitting the Bayesian observer model to psychophysical data ("behavioral prior") to the point that the two priors are indistinguishable in a cross-validation model comparison. Our results demonstrate a quantitative validation of the Bayesian observer model constrained by efficient coding at both the behavioral and neural levels.SIGNIFICANCE STATEMENT Statistical regularities of the environment play an important role in shaping both neural representations and perceptual behavior. Most previous work addressed these two aspects independently. Here we present a quantitative validation of a theoretical framework that makes joint predictions for neural coding and behavior, based on the assumption that neural representations of sensory information are efficient but also optimally used in generating a percept. Specifically, we demonstrate that the neural tuning characteristics for visual speed in brain area MT are precisely predicted by the statistical prior expectations extracted from psychophysical data. As such, our results provide a normative link between perceptual behavior and the neural representation of sensory information in the brain.

Categorical judgments do not modify sensory representations in working memory.

Categorical judgments can systematically bias the perceptual interpretation of stimulus features. However, it remained unclear whether categorical judgments directly modify working memory representations or, alternatively, generate these biases via an inference process down-stream from working memory. To address this question we ran two novel psychophysical experiments in which human subjects had to reverse their categorical judgments about a stimulus feature, if incorrect, before providing an estimate of the feature. If categorical judgments indeed directly altered sensory representations in working memory, subjects' estimates should reflect some aspects of their initial (incorrect) categorical judgment in those trials. We found no traces of the initial categorical judgment. Rather, subjects seemed to be able to flexibly switch their categorical judgment if needed and use the correct corresponding categorical prior to properly perform feature inference. A cross-validated model comparison also revealed that feedback may lead to selective memory recall such that only memory samples that are consistent with the categorical judgment are accepted for the inference process. Our results suggest that categorical judgments do not modify sensory information in working memory but rather act as top-down expectations in the subsequent sensory recall and inference process.

Individuals with autism spectrum disorder have altered visual encoding capacity.

Perceptual anomalies in individuals with autism spectrum disorder (ASD) have been attributed to an imbalance in weighting incoming sensory evidence with prior knowledge when interpreting sensory information. Here, we show that sensory encoding and how it adapts to changing stimulus statistics during feedback also characteristically differs between neurotypical and ASD groups. In a visual orientation estimation task, we extracted the accuracy of sensory encoding from psychophysical data by using an information theoretic measure. Initially, sensory representations in both groups reflected the statistics of visual orientations in natural scenes, but encoding capacity was overall lower in the ASD group. Exposure to an artificial (i.e., uniform) distribution of visual orientations coupled with performance feedback altered the sensory representations of the neurotypical group toward the novel experimental statistics, while also increasing their total encoding capacity. In contrast, neither total encoding capacity nor its allocation significantly changed in the ASD group. Across both groups, the degree of adaptation was correlated with participants' initial encoding capacity. These findings highlight substantial deficits in sensory encoding-independent from and potentially in addition to deficits in decoding-in individuals with ASD.

Benefits of commitment in hierarchical inference.

Humans have the tendency to commit to a single interpretation of what has caused some observed evidence rather than considering all possible alternatives. This tendency can explain various forms of biases in cognition and perception. However, committing to a single high-level interpretation seems short-sighted and irrational, and thus it is unclear why humans are motivated to use such strategy. In a first step toward answering this question, we systematically quantified how this strategy affects estimation accuracy at the feature level in the context of 2 common hierarchical inference tasks, category-based perception and causal cue combination. Using model simulations, we demonstrate that although estimation accuracy is generally impaired when conditioned on only a single high-level interpretation, the reduction is not uniform across the entire feature range. Compared with a full inference strategy that considers all high-level interpretations, accuracy is only worse for feature values relatively close to the decision boundaries but is better everywhere else. That is, for feature values for which an observer has a reasonably high chance of being correct about the high-level interpretation of the feature, a full commitment to that particular interpretation is advantageous. We also show that conditioning on an preceding high-level interpretation provides an effective mechanism for partially protecting the evidence from corruption with late noise in the inference process (e.g., during retention in and recall from working memory). Our results suggest that a top-down inference strategy that solely relies on the most likely high-level interpretation can be favorable with regard to late noise and more holistic performance metrics. (PsycInfo Database Record (c) 2020 APA, all rights reserved).

Post-decision biases reveal a self-consistency principle in perceptual inference.

Making a categorical judgment can systematically bias our subsequent perception of the world. We show that these biases are well explained by a self-consistent Bayesian observer whose perceptual inference process is causally conditioned on the preceding choice. We quantitatively validated the model and its key assumptions with a targeted set of three psychophysical experiments, focusing on a task sequence where subjects first had to make a categorical orientation judgment before estimating the actual orientation of a visual stimulus. Subjects exhibited a high degree of consistency between categorical judgment and estimate, which is difficult to reconcile with alternative models in the face of late, memory related noise. The observed bias patterns resemble the well-known changes in subjective preferences associated with cognitive dissonance, which suggests that the brain's inference processes may be governed by a universal self-consistency constraint that avoids entertaining 'dissonant' interpretations of the evidence.

Credo for optimality.

Optimal or suboptimal, Rahnev & Denison (R&D) rightly argue that this ill-defined distinction is not useful when comparing models of perceptual decision making. However, what they miss is how valuable the focus on optimality has been in deriving these models in the first place. Rather than prematurely abandon the optimality assumption, we should refine this successful normative hypothesis with additional constraints that capture specific limitations of (sensory) information processing in the brain.

Lawful relation between perceptual bias and discriminability.

Perception of a stimulus can be characterized by two fundamental psychophysical measures: how well the stimulus can be discriminated from similar ones (discrimination threshold) and how strongly the perceived stimulus value deviates on average from the true stimulus value (perceptual bias). We demonstrate that perceptual bias and discriminability, as functions of the stimulus value, follow a surprisingly simple mathematical relation. The relation, which is derived from a theory combining optimal encoding and decoding, is well supported by a wide range of reported psychophysical data including perceptual changes induced by contextual modulation. The large empirical support indicates that the proposed relation may represent a psychophysical law in human perception. Our results imply that the computational processes of sensory encoding and perceptual decoding are matched and optimized based on identical assumptions about the statistical structure of the sensory environment.

Visual Decision-Making in an Uncertain and Dynamic World.

The right decision today may be the wrong decision tomorrow. We live in a world in which expectations, contingencies, and goals continually evolve and change. Thus, decisions do not occur in isolation but rather are tightly embedded in these streams of temporal dependencies. Accordingly, even relatively straightforward visual decisions must take into account not just the immediate sensory input but also past experiences and future goals and expectations. Here, we evaluate recent progress in understanding how the brain implements these dependencies. We show that visual decision-making relies on mechanisms of evidence accumulation and commitment that have been studied extensively under relatively static, isolated conditions but in general can operate much more flexibly. A deeper understanding of these mechanisms will require identifying the principles that govern this flexibility, which must operate across different timescales to produce effective decisions in uncertain and dynamic environments.

Efficient Neural Codes That Minimize Lp Reconstruction Error.

The efficient coding hypothesis assumes that biological sensory systems use neural codes that are optimized to best possibly represent the stimuli that occur in their environment. Most common models use information-theoretic measures, whereas alternative formulations propose incorporating downstream decoding performance. Here we provide a systematic evaluation of different optimality criteria using a parametric formulation of the efficient coding problem based on the [Formula: see text] reconstruction error of the maximum likelihood decoder. This parametric family includes both the information maximization criterion and squared decoding error as special cases. We analytically derived the optimal tuning curve of a single neuron encoding a one-dimensional stimulus with an arbitrary input distribution. We show how the result can be generalized to a class of neural populations by introducing the concept of a meta-tuning curve. The predictions of our framework are tested against previously measured characteristics of some early visual systems found in biology. We find solutions that correspond to low values of [Formula: see text], suggesting that across different animal models, neural representations in the early visual pathways optimize similar criteria about natural stimuli that are relatively close to the information maximization criterion.

Mutual Information, Fisher Information, and Efficient Coding.

Fisher information is generally believed to represent a lower bound on mutual information (Brunel & Nadal, 1998), a result that is frequently used in the assessment of neural coding efficiency. However, we demonstrate that the relation between these two quantities is more nuanced than previously thought. For example, we find that in the small noise regime, Fisher information actually provides an upper bound on mutual information. Generally our results show that it is more appropriate to consider Fisher information as an approximation rather than a bound on mutual information. We analytically derive the correspondence between the two quantities and the conditions under which the approximation is good. Our results have implications for neural coding theories and the link between neural population coding and psychophysically measurable behavior. Specifically, they allow us to formulate the efficient coding problem of maximizing mutual information between a stimulus variable and the response of a neural population in terms of Fisher information. We derive a signature of efficient coding expressed as the correspondence between the population Fisher information and the distribution of the stimulus variable. The signature is more general than previously proposed solutions that rely on specific assumptions about the neural tuning characteristics. We demonstrate that it can explain measured tuning characteristics of cortical neural populations that do not agree with previous models of efficient coding.

A Bayesian observer model constrained by efficient coding can explain 'anti-Bayesian' percepts.

Bayesian observer models provide a principled account of the fact that our perception of the world rarely matches physical reality. The standard explanation is that our percepts are biased toward our prior beliefs. However, reported psychophysical data suggest that this view may be simplistic. We propose a new model formulation based on efficient coding that is fully specified for any given natural stimulus distribution. The model makes two new and seemingly anti-Bayesian predictions. First, it predicts that perception is often biased away from an observer's prior beliefs. Second, it predicts that stimulus uncertainty differentially affects perceptual bias depending on whether the uncertainty is induced by internal or external noise. We found that both model predictions match reported perceptual biases in perceived visual orientation and spatial frequency, and were able to explain data that have not been explained before. The model is general and should prove applicable to other perceptual variables and tasks.

Signal Integration in Human Visual Speed Perception.

Object motion in natural scenes results in visual stimuli with a rich and broad spatiotemporal frequency spectrum. While the question of how the visual system detects and senses motion energies at different spatial and temporal frequencies has been fairly well studied, it is unclear how the visual system integrates this information to form coherent percepts of object motion. We applied a combination of tailored psychophysical experiments and predictive modeling to address this question with regard to perceived motion in a given direction (i.e., stimulus speed). We tested human subjects in a discrimination experiment using stimuli that selectively targeted four distinct spatiotemporally tuned channels with center frequencies consistent with a common speed. We first characterized subjects' responses to stimuli that targeted only individual channels. Based on these measurements, we then predicted subjects' psychometric functions for stimuli that targeted multiple channels simultaneously. Specifically, we compared predictions of three Bayesian observer models that either optimally integrated the information across all spatiotemporal channels, or only used information from the most reliable channel, or formed an average percept across channels. Only the model with optimal integration was successful in accounting for the data. Furthermore, the proposed channel model provides an intuitive explanation for the previously reported spatial frequency dependence of perceived speed of coherent object motion. Finally, our findings indicate that a prior expectation for slow speeds is added to the inference process only after the sensory information is combined and integrated.

Characterizing the impact of category uncertainty on human auditory categorization behavior.

Categorization is an important cognitive process. However, the correct categorization of a stimulus is often challenging because categories can have overlapping boundaries. Whereas perceptual categorization has been extensively studied in vision, the analogous phenomenon in audition has yet to be systematically explored. Here, we test whether and how human subjects learn to use category distributions and prior probabilities, as well as whether subjects employ an optimal decision strategy when making auditory-category decisions. We asked subjects to classify the frequency of a tone burst into one of two overlapping, uniform categories according to the perceived tone frequency. We systematically varied the prior probability of presenting a tone burst with a frequency originating from one versus the other category. Most subjects learned these changes in prior probabilities early in testing and used this information to influence categorization. We also measured each subject's frequency-discrimination thresholds (i.e., their sensory uncertainty levels). We tested each subject's average behavior against variations of a Bayesian model that either led to optimal or sub-optimal decision behavior (i.e. probability matching). In both predicting and fitting each subject's average behavior, we found that probability matching provided a better account of human decision behavior. The model fits confirmed that subjects were able to learn category prior probabilities and approximate forms of the category distributions. Finally, we systematically explored the potential ways that additional noise sources could influence categorization behavior. We found that an optimal decision strategy can produce probability-matching behavior if it utilized non-stationary category distributions and prior probabilities formed over a short stimulus history. Our work extends previous findings into the auditory domain and reformulates the issue of categorization in a manner that can help to interpret the results of previous research within a generative framework.

A new two-alternative forced choice method for the unbiased characterization of perceptual bias and discriminability.

Perception is often biased by secondary stimulus attributes (e.g., stimulus noise, attention, or spatial context). A correct quantitative characterization of perceptual bias is essential for testing hypotheses about the underlying perceptual mechanisms and computations. We demonstrate that the standard two-alternative forced choice (2AFC) method can lead to incorrect estimates of perceptual bias. We present a new 2AFC method that solves this problem by asking subjects to judge the relative perceptual distances between the test and each of two reference stimuli. Naïve subjects can easily perform this task. We successfully validated the new method with a visual motion-discrimination experiment. We demonstrate that the method permits an efficient and accurate characterization of perceptual bias and simultaneously provides measures of discriminability for both the reference and test stimulus, all from a single stimulus condition. This makes it an attractive choice for the characterization of perceptual bias and discriminability in a wide variety of psychophysical experiments.

Dynamic belief state representations.

Perceptual and control systems are tasked with the challenge of accurately and efficiently estimating the dynamic states of objects in the environment. To properly account for uncertainty, it is necessary to maintain a dynamical belief state representation rather than a single state vector. In this review, canonical algorithms for computing and updating belief states in robotic applications are delineated, and connections to biological systems are highlighted. A navigation example is used to illustrate the importance of properly accounting for correlations between belief state components, and to motivate the need for further investigations in psychophysics and neurobiology.