Biology subjects

Ma, W. J.

Publications and source records attributed to Ma, W. J..

14 recordsLinked to original sources

bioRxiv2018

A neural basis of probabilistic computation in visual cortex

Bayesian models of behavior suggest that organisms represent uncertainty associated with sensory variables. However, the neural code of uncertainty remains elusive. A central hypothesis is that uncertainty is encoded in the population activity of cortical neurons in the form of likelihood functions. We studied the neural code of uncertainty by simultaneously recording population activity from the primate visual cortex during a visual categorization task in which trial-to-trial uncertainty about stimulus orientation was relevant for the decision. We decoded the likelihood function from the trial-to-trial population activity and found that it predicted decisions better than a point estimate of orientation. This remained true when we conditioned on the true orientation, suggesting that internal fluctuations in neural activity drive behaviorally meaningful variations in the likelihood function. Our results establish the role of population-encoded likelihood functions in mediating behavior, and provide a neural underpinning for Bayesian models of perception.

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bioRxiv2018

Sources of suboptimality in a minimalistic explore-exploit task

Balancing exploration and exploitation is a fundamental aspect of decision-making. It remains unknown whether people are close to optimal in striking this balance, and if not, how exactly their behavior deviates from optimality. Many existing paradigms are not ideally suited to answer this question, as they contain complexities such as non-stationary environments, stochasticity under exploitation, and reward distributions that are unknown to participants. Here, we introduce a task without such complexities, in which the optimal policy is to start off exploring and to switch to exploitation at most once in each sequence of decisions. The behavior of 49 laboratory and 143 online participants deviated both qualitatively and quantitatively from the optimal policy, even when allowing for bias and decision noise. Instead, people seem to follow a suboptimal rule in which they switch from exploration to exploitation when the highest reward so far exceeds a certain threshold. Moreover, we show that this threshold decreases approximately linearly with the proportion of the sequence that remains, suggesting a novel temporal ratio law. Finally, we find evidence for \"sequence-level\" variability which is shared across all decisions in the same sequence. Our results provide a new perspective on the explore-exploit dilemma, and emphasize the importance of examining sequence-level strategies and their variability when studying sequential decision-making.

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bioRxiv2018

The Role of Sensory Uncertainty in Simple Perceptual Organization

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 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. Author summaryOur percept of the world is governed not only by the sensory information we have access to, but also by the way we interpret this information. When presented with a visual scene, our visual system undergoes a process of grouping visual elements together to form coherent entities so that we can interpret the scene more readily and meaningfully. For example, when looking at a pile of autumn leaves, one can still perceive and identify a whole leaf even when it is partially covered by another leaf. While Gestalt psychologists have long described perceptual organization with a set of qualitative laws, recent studies offered a statistically-optimal - Bayesian, in statistical jargon - interpretation of this process, whereby the observer chooses the scene configuration with the highest probability given the available sensory inputs. However, these studies drew their conclusions without considering a key actor in this kind of statistically-optimal computations, that is the role of sensory uncertainty. One can easily imagine that our decision on whether two contours belong to the same leaf or different leaves is likely going to change when we move from viewing the pile of leaves at a great distance (high sensory uncertainty), to viewing very closely (low sensory uncertainty). Our study examines whether and how people incorporate uncertainty into contour integration, an elementary form of perceptual organization, by varying sensory uncertainty from trial to trial in a simple contour integration task. We found that people indeed take into account sensory uncertainty, however in a way that subtly deviates from optimal behavior.

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bioRxiv2018

Strategic allocation of working memory resource

Visual working memory (VWM), the brief retention of past visual information, supports a range of cognitive functions (Fukuda, Vogel, Mayr, & Awh, 2010; Johnson et al., 2013). The resource that supports VWM is limited, raising the question of how the brain allocates this limited resource to different objects. This question is even more interesting in ecological settings, in which objects are not equally important. In a psychophysical experiment, participants remembered the location of four targets with different probabilities of being tested after a delay. We then measured their memory accuracy of one of the targets. We found that participants allocated more resource to memoranda with higher priority, but underallocated resource to high- and overallocated to low-priority targets relative to the true probe probabilities. These results are well explained by a computational model in which resource is allocated to minimize expected estimation error. We replicated this finding in a second experiment in which participants bet on their memory fidelity after making the location estimate. The results of this experiment show that people 1) use information about memory quality and 2) minimize error even with an incentivized, alternative resource allocation strategy. Humans may mitigate the behavioral effects of a limited VWM through knowledge of memory fidelity and strategic resource allocation.

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bioRxiv2018

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

In daily life, working memory 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 is poorly understood. To investigate this, we conducted a memory experiment where people not only reported an estimate of a remembered stimulus, but also made a rewarded decision designed to reflect memory uncertainty. Reported memory uncertainty is correlated with estimation error, showing people incorporate their trial-to-trial memory quality into rewarded decisions. Moreover, memory uncertainty can be combined with other sources of information; after we induced prior beliefs about stimuli probabilities, we found that estimates shifted towards more frequent colors, with the shift increasing with reported uncertainty. The data is best predicted by models where people incorporate their trial-to-trial memory uncertainty with potential rewards and prior beliefs, highlighting the importance of studying working memory as a process integrated with decision-making.

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bioRxiv2018

Problematic usage of the Zhang and Luck mixture model

A common method, due to Zhang and Luck (2008), for analyzing delayed-estimation data with a circular stimulus variable is to fit a mixture of a Von Mises distribution and a uniform distribution. The uniform distribution represents random guesses, presumably made when an item is not kept in memory. When I generate synthetic data from a variable-precision model with zero guessing, the method estimates the guess rate to be nonzero and often high. This is due to model mismatch: the fitted model is not matched to the data-generating (true) model. In real data, this could be a problem if one considers the variable-precision model a plausible candidate model and draws conclusions based on the estimated guess rates. I describe five solutions to this problem: analyzing the residual, ruling out the variable-precision model, robust inference, fitting a hybrid model, and using model-free statistics. I hope that these solutions can contribute to good data analysis practices in the study of working memory.

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bioRxiv2018

A Diverse Range of Factors Affect the Nature of Neural Representations Underlying Short-Term Memory

Sequential and persistent activity models are two prominent models of short-term memory in neural circuits. In persistent activity models, memories are represented in persistent or nearly persistent activity patterns across a population of neurons, whereas in sequential models, memories are represented dynamically by a sequential pattern of activity across the population. Experimental evidence for both types of model in the brain has been reported previously. However, it has been unclear under what conditions these two qualitatively different types of solutions emerge in neural circuits. Here, we address this question by training recurrent neural networks on several short-term memory tasks under a wide range of circuit and task manipulations. We show that sequential and nearly persistent solutions are both part of a spectrum that emerges naturally in trained networks under different conditions. Fixed delay durations, tasks with higher temporal complexity, strong network coupling, motion-related dynamic inputs and prior training in a different task favor more sequential solutions, whereas variable delay durations, tasks with low temporal complexity, weak network coupling and symmetric Hebbian short-term synaptic plasticity favor more persistent solutions. Our results help clarify some seemingly contradictory experimental results on the existence of sequential vs. persistent activity based memory mechanisms in the brain.

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bioRxiv2017

Limitations of proposed signatures of Bayesian confidence

The Bayesian model of confidence posits that confidence is the observers posterior probability that the decision is correct. It has been proposed that researchers can gain evidence in favor of the Bayesian model by deriving qualitative signatures of Bayesian confidence, i.e., patterns that one would expect to see if an observer was Bayesian, and looking for those signatures in human or animal data. We examine two proposed qualitative 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 for trials with neutral evidence. This signature only holds when class-conditioned stimulus distributions do not overlap and internal noise is very low. Another signature is that, as stimulus magnitude increases, confidence increases on correct trials but decreases on incorrect trials. This signature is also dependent on stimulus distribution type. There is an alternative form of this signature that has been applied in the literature; we find no indication that it is expected under Bayesian confidence, which resolves an ostensible discrepancy. We conclude that, to determine the nature of the computations underlying confidence reports, there may be no shortcut to quantitative model comparison.

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bioRxiv2017

A low-level perceptual correlate of behavioral and clinical deficits in ADHD

In many studies of attention-deficit hyperactivity disorder (ADHD), stimulus encoding and processing (per-ceptual function) and response selection (executive function) have been intertwined. To dissociate deficits in these functions, we introduced a task that parametrically varied low-level stimulus features (orientation and color) for fine-grained analysis of perceptual function. It also required participants to switch their attention between feature dimensions on a trial-by-trial basis, thus taxing executive processes. Furthermore, we used a response paradigm that captured task-irrelevant motor output (TIMO), reflecting failures to use the correct stimulus-response rule. ADHD participants had substantially higher perceptual variability than Controls, especially for orientation, as well as higher TIMO. In both ADHD and Controls, TIMO was strongly affected by the switch manipulation. Across participants, the perceptual variability parameter was correlated with TIMO, suggesting that perceptual deficits are associated with executive function deficits. Based on perceptual variability alone, we were able to classify participants into ADHD and Controls with a mean accuracy of about 77%. Participants self-reported General Executive Composite score correlated not only with TIMO but also with the perceptual variability parameter. Our results highlight the role of perceptual deficits in ADHD and the usefulness of computational modeling of behavior in dissociating perceptual from executive processes.

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bioRxiv2017

Humans flexibly 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 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 observers attentional state on each trial contributed probabilistically to the decision computation. This responsiveness of an observers decisions to attention-dependent uncertainty should improve perceptual decisions in natural vision, in which attention is unevenly distributed across a scene.

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bioRxiv2017

Quantifying the importance of guessing, decision noise, and variable precision in explaining behavioral variability in visual perception

Given the same sensory stimuli in the same task, human observers do not always make the same response. Well-known sources of behavioral variability are sensory noise and guessing. Visual short-term memory studies have suggested that the precision of the sensory noise is itself variable. However, it is unknown whether precision is also variable in perceptual tasks without a memory component. We searched for evidence for variable precision in 11 visual perception tasks with a single relevant feature, orientation. We specifically examined the effect of distractor stimuli: distractors were absent, homogeneous and fixed across trials, homogeneous and variable, or heterogeneous and variable. We first considered four models: with and without guessing, and with and without variability in precision. We quantified the importance of both factors using six metrics: factor knock-in difference, factor knock-out difference, and log factor posterior ratio, each based on AIC or BIC. According to all six metrics, we found strong evidence for variable precision in five experiments. Next, we extended our model space to include potential confounding factors: the oblique effect and decision noise. This left strong evidence for variable precision in only one experiment, in which distractors were homogeneous but variable. Finally, when we considered suboptimal decision rules, the evidence also disappeared in this experiment. Our results provide little evidence for variable precision overall and only a hint when distractors are variable. Methodologically, the results underline the importance of including multiple factors in factorial model comparison: testing for only two factors would have yielded an incorrect conclusion.

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bioRxiv2017

A rational theory of the limitations of working memory and attention

Encoding precision in visual working memory decreases with the number of encoded items. Here, we propose a normative theory for such set size effects: the brain minimizes a weighted sum of an error-based behavioral cost and a neural encoding cost. We construct a model from this theory and find that it predicts set size effects. Notably, these effects are mediated by probing probability, which aligns with previous empirical findings. The model accounts well for effects of both set size and probing probability on encoding precision in nine delayed-estimation experiments. Moreover, we find support for the prediction that the total amount of invested resource can vary non-monotonically with set size. Finally, we show that it is sometimes optimal to encode only a subset or even none of the relevant items in a task. Our findings raise the possibility that cognitive \"limitations\" arise from rational cost minimization rather than from constraints.

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bioRxiv2017

Bayesian Comparison of Explicit and Implicit Causal Inference Strategies in Multisensory Heading Perception

The precision of multisensory heading perception improves when visual and vestibular cues arising from the same cause, namely motion of the observer through a stationary environment, are integrated. Thus, 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 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.

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bioRxiv2016

The computations underlying human confidence reports are probabilistic, but not Bayesian

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. However, behavior is not fully consistent with the Bayesian hypothesis and is better described by simple heuristic models. 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.

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