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<front>
<journal-meta>
<journal-id journal-id-type="nlm-ta">PLoS ONE</journal-id>
<journal-id journal-id-type="publisher-id">plos</journal-id>
<journal-id journal-id-type="pmc">plosone</journal-id><journal-title-group>
<journal-title>PLoS ONE</journal-title></journal-title-group>
<issn pub-type="epub">1932-6203</issn>
<publisher>
<publisher-name>Public Library of Science</publisher-name>
<publisher-loc>San Francisco, USA</publisher-loc></publisher>
</journal-meta>
<article-meta>
<article-id pub-id-type="publisher-id">PONE-D-13-52613</article-id>
<article-id pub-id-type="doi">10.1371/journal.pone.0096511</article-id>
<article-categories><subj-group subj-group-type="heading"><subject>Research Article</subject></subj-group><subj-group subj-group-type="Discipline-v2"><subject>Biology and life sciences</subject><subj-group><subject>Computational biology</subject><subj-group><subject>Computational neuroscience</subject></subj-group></subj-group><subj-group><subject>Neuroscience</subject><subj-group><subject>Cognitive science</subject><subj-group><subject>Cognition</subject><subj-group><subject>Decision making</subject></subj-group></subj-group></subj-group><subj-group><subject>Cognitive neuroscience</subject></subj-group></subj-group><subj-group><subject>Psychology</subject><subj-group><subject>Behavior</subject><subj-group><subject>Human performance</subject></subj-group></subj-group><subj-group><subject>Psychometrics</subject></subj-group></subj-group></subj-group></article-categories>
<title-group>
<article-title>Relation between Belief and Performance in Perceptual Decision Making</article-title>
<alt-title alt-title-type="running-head">Belief/Performance in Perceptual Decision Making</alt-title>
</title-group>
<contrib-group>
<contrib contrib-type="author" xlink:type="simple"><name name-style="western"><surname>Drugowitsch</surname><given-names>Jan</given-names></name><xref ref-type="aff" rid="aff1"><sup>1</sup></xref><xref ref-type="aff" rid="aff2"><sup>2</sup></xref><xref ref-type="aff" rid="aff3"><sup>3</sup></xref><xref ref-type="corresp" rid="cor1"><sup>*</sup></xref></contrib>
<contrib contrib-type="author" xlink:type="simple"><name name-style="western"><surname>Moreno-Bote</surname><given-names>Rubén</given-names></name><xref ref-type="aff" rid="aff4"><sup>4</sup></xref><xref ref-type="aff" rid="aff5"><sup>5</sup></xref></contrib>
<contrib contrib-type="author" xlink:type="simple"><name name-style="western"><surname>Pouget</surname><given-names>Alexandre</given-names></name><xref ref-type="aff" rid="aff1"><sup>1</sup></xref><xref ref-type="aff" rid="aff3"><sup>3</sup></xref></contrib>
</contrib-group>
<aff id="aff1"><label>1</label><addr-line>Department of Brain and Cognitive Sciences, University of Rochester, Rochester, New York, United States of America</addr-line></aff>
<aff id="aff2"><label>2</label><addr-line>Institut National de la Santé et de la Recherche Médicale, École Normale Supérieure, Paris, France</addr-line></aff>
<aff id="aff3"><label>3</label><addr-line>Département des Neurosciences Fondamentales, Université de Genève, Geneva, Switzerland</addr-line></aff>
<aff id="aff4"><label>4</label><addr-line>Research Unit, Parc Sanitari Sant Joan de Déu and Universitat de Barcelona, Barcelona, Spain</addr-line></aff>
<aff id="aff5"><label>5</label><addr-line>Centro de Investigación Biomédica en Red de Salud Mental (CIBERSAM), Barcelona, Spain</addr-line></aff>
<contrib-group>
<contrib contrib-type="editor" xlink:type="simple"><name name-style="western"><surname>Hamed</surname><given-names>Suliann Ben</given-names></name>
<role>Editor</role>
<xref ref-type="aff" rid="edit1"/></contrib>
</contrib-group>
<aff id="edit1"><addr-line>Centre de Neuroscience Cognitive, France</addr-line></aff>
<author-notes>
<corresp id="cor1">* E-mail: <email xlink:type="simple">jdrugo@gmail.com</email></corresp>
<fn fn-type="conflict"><p>The authors have declared that no competing interests exist.</p></fn>
<fn fn-type="con"><p>Conceived and designed the experiments: JD RMB AP. Performed the experiments: JD RMB. Analyzed the data: JD. Wrote the paper: JD RMB AP.</p></fn>
</author-notes>
<pub-date pub-type="collection"><year>2014</year></pub-date>
<pub-date pub-type="epub"><day>9</day><month>5</month><year>2014</year></pub-date>
<volume>9</volume>
<issue>5</issue>
<elocation-id>e96511</elocation-id>
<history>
<date date-type="received"><day>26</day><month>12</month><year>2013</year></date>
<date date-type="accepted"><day>8</day><month>4</month><year>2014</year></date>
</history>
<permissions>
<copyright-year>2014</copyright-year>
<copyright-holder>Drugowitsch et al</copyright-holder><license xlink:type="simple"><license-p>This is an open-access article distributed under the terms of the <ext-link ext-link-type="uri" xlink:href="http://creativecommons.org/licenses/by/4.0/" xlink:type="simple">Creative Commons Attribution License</ext-link>, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.</license-p></license></permissions>
<abstract>
<p>In an uncertain and ambiguous world, effective decision making requires that subjects form and maintain a belief about the correctness of their choices, a process called meta-cognition. Prediction of future outcomes and self-monitoring are only effective if belief closely matches behavioral performance. Equality between belief and performance is also critical for experimentalists to gain insight into the subjects' belief by simply measuring their performance. Assuming that the decision maker holds the correct model of the world, one might indeed expect that belief and performance should go hand in hand. Unfortunately, we show here that this is rarely the case when performance is defined as the percentage of correct responses for a fixed stimulus, a standard definition in psychophysics. In this case, belief equals performance only for a very narrow family of tasks, whereas in others they will only be very weakly correlated. As we will see it is possible to restore this equality in specific circumstances but this remedy is only effective for a decision-maker, not for an experimenter. We furthermore show that belief and performance do not match when conditioned on task difficulty – as is common practice when plotting the psychometric curve – highlighting common pitfalls in previous neuroscience work. Finally, we demonstrate that miscalibration and the hard-easy effect observed in humans' and other animals' certainty judgments could be explained by a mismatch between the experimenter's and decision maker's expected distribution of task difficulties. These results have important implications for experimental design and are of relevance for theories that aim to unravel the nature of meta-cognition.</p>
</abstract>
<funding-group><funding-statement>R.M.B. was supported by a Ramón y Cajal Spanish Award and by the Marie Curie FP7-PEOPLE-2010-IRG grant (PIRG08-GA-2010-276795). A.P. was supported by grants from the National Science Foundation (BCS0446730 and REESE 1109366), a Multidisciplinary University Research Initiative (N00014-07-1-0937) and the James McDonnell Foundation. The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript.</funding-statement></funding-group><counts><page-count count="16"/></counts></article-meta>
</front>
<body><sec id="s1">
<title>Introduction</title>
<p>In an uncertain and ambiguous world, effective decision making requires computing one's certainty about all decision-relevant evidence. For example, consider driving on the highway while running late for a job interview. Driving too fast would result in a very high cost if hit by another car. Driving too slowly, on the other hand, could result in losing the job. Thus, a good policy to follow is to accumulate evidence about the surrounding traffic to minimize the expected personal cost of an accident, evaluated based on ones certainty, while balancing the loss of time to accumulate this evidence. In general, decision certainty plays an essential role in value-based decisions, and is thus an essential component of every-day decision making. There exists a large body of evidence that humans and animals encode such information, which allows them to feature a belief, or confidence, about the correctness of their decisions (a process sometimes referred to as meta-cognition) <xref ref-type="bibr" rid="pone.0096511-Beck1">[1]</xref>–<xref ref-type="bibr" rid="pone.0096511-Smith1">[10]</xref>. It is important to mention that in this paper it is not claimed that belief is explicit, conscious or readily accessible for verbal report. Rather, belief can be implicitly coded (e.g., a function of several variables of the decision process), unconscious in many cases and difficult - if not impossible - to access verbally.</p>
<p>Nevertheless, for the decision maker, such belief is important as predicting the decision's outcome and monitoring her task performance are only effective if this belief is correctly reflected in the decision maker's performance. The relation between belief and performance is also essential for an experimenter who wants to assess the decision maker's belief to gain insight into her decision making strategy <xref ref-type="bibr" rid="pone.0096511-Kiani1">[5]</xref>, <xref ref-type="bibr" rid="pone.0096511-Kepecs1">[8]</xref> by, for example, using the decision maker's performance as a proxy for her belief <xref ref-type="bibr" rid="pone.0096511-Grinband1">[11]</xref>. In both cases, belief and performance are assumed to be closely related or equivalent.</p>
<p>Assuming that the decision maker holds the correct model of the world, it is intuitive that her belief should equal her performance <xref ref-type="bibr" rid="pone.0096511-Gigerenzer1">[12]</xref>. For instance, if a subject is correct 80% of the time across trials of a particular experimental condition, it seems logical to conclude that, on any given trial, the subjects should believe that her chances of being correct is 80%. Indeed, some previous studies on decision making have implicitly assumed these measures to be similar <xref ref-type="bibr" rid="pone.0096511-Kiani1">[5]</xref>, <xref ref-type="bibr" rid="pone.0096511-Kepecs1">[8]</xref> or even exchangeable <xref ref-type="bibr" rid="pone.0096511-Grinband1">[11]</xref>. Surprisingly, however, we show that belief equals performance only for a very narrow family of tasks and decision strategies. So, if a subject has the correct model of the world, how is it possible that her belief does not correspond to her performance in most realistic conditions? And if that is the case, how can subjects trust their belief to monitor their performance in order to improve it in any given task?</p>
<p>The theory that we outline below reveals (i) the correct variables that a decision maker should monitor during a task, (ii) the conditions under which an experimenter (that is, and external observer controlling some variables of the task at hand) can measure belief at each trial or on average, and (iii) the correct performance measures to be used to estimate the decision maker's belief without bias and with the least possible variance. Our theory is based on a normative view of the decision-making process, in which the decision maker utilizes the correct model of the world to infer optimal decisions given all available evidence. To this respect, our approach differs from comparable, but heuristic explanations for human and animal confidence judgments <xref ref-type="bibr" rid="pone.0096511-Kepecs1">[8]</xref>, <xref ref-type="bibr" rid="pone.0096511-Ratcliff1">[13]</xref>–<xref ref-type="bibr" rid="pone.0096511-Pleskac1">[15]</xref> that might employ comparable mechanisms but do not have the same ideological underpinning. As such, our theory provides an upper bound on the relation between belief and performance. Despite this, we demonstrate some significant deterioration of this relation, which can, due to deviations from the normative ideal, only worsen in practice. Based on these findings, we point out some pitfalls in previous neuroscience work, we provide a new hypothesis for the origin of the hard-easy effect, and we present a different perspective on models of confidence miscalibration <xref ref-type="bibr" rid="pone.0096511-Juslin1">[16]</xref>–<xref ref-type="bibr" rid="pone.0096511-Moore1">[18]</xref>.</p>
<p>We first introduce the general formalism, based upon which we define belief and performance. This is followed by discussing their relation and showing that they are rarely equivalent. We then focus on the more specific case of diffusion and race decision making models, and demonstrate how our general findings apply to these two model types. After that, we discuss the consequences of these findings to both the decision-maker and the experimenter observing this decision-maker, focusing on the relation between the psychometric curve and the decision maker's belief, and the hard-easy effect in human confidence judgments. At last, we put our findings into the more general context of previous work.</p>
</sec><sec id="s2">
<title>Results</title>
<sec id="s2a">
<title>Formalism</title>
<p>In general, we consider <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e001" xlink:type="simple"/></inline-formula>-alternative forced choice (<inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e002" xlink:type="simple"/></inline-formula>-AFC) tasks (<inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e003" xlink:type="simple"/></inline-formula>) with a sequence of independent trials, in each of which an experimenter determines the hidden state <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e004" xlink:type="simple"/></inline-formula> of the world, and the aim of the decision maker is to identify this state based on limited information (<xref ref-type="fig" rid="pone-0096511-g001">Fig. 1</xref>). At the beginning of each trial, the experimenter draws the hidden state <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e005" xlink:type="simple"/></inline-formula> from the prior probability distribution <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e006" xlink:type="simple"/></inline-formula>. This state can take one of <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e007" xlink:type="simple"/></inline-formula> values out of the set <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e008" xlink:type="simple"/></inline-formula>. Consider, for example, an orientation categorization task, in which a displayed orientation is generated stochastically from one of two categories, and the decision maker's task is to identify this category upon observing the orientation. In this example, we would have <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e009" xlink:type="simple"/></inline-formula>, such that the generative category <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e010" xlink:type="simple"/></inline-formula> can take values out of the set <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e011" xlink:type="simple"/></inline-formula>. Furthermore, if each category is a-priory equally likely, we would have <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e012" xlink:type="simple"/></inline-formula>.</p>
<fig id="pone-0096511-g001" position="float"><object-id pub-id-type="doi">10.1371/journal.pone.0096511.g001</object-id><label>Figure 1</label><caption>
<title>Illustration of framework with a three-sided coin example.</title>
<p>(a) In each trial of a sequence, a hidden state <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e013" xlink:type="simple"/></inline-formula> is picked by the experimenter, based on which the observation <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e014" xlink:type="simple"/></inline-formula> is generated. The decision maker only observes <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e015" xlink:type="simple"/></inline-formula> but not <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e016" xlink:type="simple"/></inline-formula> and chooses option <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e017" xlink:type="simple"/></inline-formula> where <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e018" xlink:type="simple"/></inline-formula> is a deterministic function that maps observations into decisions. In this 2-AFC example there are two possible hidden state, causing <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e019" xlink:type="simple"/></inline-formula> to be sampled either according to a biased 3-sided coin <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e020" xlink:type="simple"/></inline-formula>, or a fair 3-sided coin <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e021" xlink:type="simple"/></inline-formula>. (b) For the given decision function, which maximizes the number of correct decisions for <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e022" xlink:type="simple"/></inline-formula> and <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e023" xlink:type="simple"/></inline-formula>, the resulting belief and performance are shown for either choice/hidden state. Belief and performance only match if <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e024" xlink:type="simple"/></inline-formula>, that is, when <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e025" xlink:type="simple"/></inline-formula>.</p>
</caption><graphic mimetype="image" xlink:href="info:doi/10.1371/journal.pone.0096511.g001" position="float" xlink:type="simple"/></fig>
<p>The decision maker does not have direct access to the hidden state <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e026" xlink:type="simple"/></inline-formula>, but instead observes some <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e027" xlink:type="simple"/></inline-formula> (for example, the displayed orientation) that is stochastically related to <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e028" xlink:type="simple"/></inline-formula> by the generative model <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e029" xlink:type="simple"/></inline-formula> (how the experimenter generates orientations for each category). Based on the observation <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e030" xlink:type="simple"/></inline-formula>, which might represent sensory input (the image of the displayed orientation on a screen) or neural activity (the firing rate of orientation-selective neurons in area V1), the decision maker commits to the choice <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e031" xlink:type="simple"/></inline-formula> by utilizing the deterministic decision function <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e032" xlink:type="simple"/></inline-formula> (we will write <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e033" xlink:type="simple"/></inline-formula> whenever we need to be explicit about its relation to <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e034" xlink:type="simple"/></inline-formula>). Thus, we assume that all stochasticity from the decision maker's choices has its origin in the stochasticity of how observations are generated from the hidden state (but see Generalizations). In that sense, what we called <italic>observation</italic> is similar to the <italic>decision variable</italic> in Signal Detection Theory <xref ref-type="bibr" rid="pone.0096511-Green1">[19]</xref>, and our decision function <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e035" xlink:type="simple"/></inline-formula> is a generalized version of the threshold that the decision variable is compared to. In addition to a deterministic decision function, we assume that the decision maker knows (for example, through experience) both the prior <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e036" xlink:type="simple"/></inline-formula> and the generative model <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e037" xlink:type="simple"/></inline-formula>, such that she could, for example, employ the decision function <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e038" xlink:type="simple"/></inline-formula> that maximizes her posterior belief <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e039" xlink:type="simple"/></inline-formula>. In our orientation categorization example, this would correspond to choosing always the category that was the most likely to have generated the observed orientation. While this might be a sensible function to use in general, our exposition is also valid for any other arbitrary choice of the decision function.</p>
<p>We will consider situations in which the experimenter has no or only limited access to the observation <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e040" xlink:type="simple"/></inline-formula> as perceived by the decision maker. For example, <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e041" xlink:type="simple"/></inline-formula> might represent the decision maker's neural activity in response to the displayed orientation, and the experimenter only observes the decision maker's choices, as determined by <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e042" xlink:type="simple"/></inline-formula>. One could also imagine that the experimenter only has control over the generative category, is unable to observe the stimulus orientations in individual trials. In both cases, the experimenter cannot know <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e043" xlink:type="simple"/></inline-formula> with certainty as many different values of <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e044" xlink:type="simple"/></inline-formula>could lead to the same decision <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e045" xlink:type="simple"/></inline-formula>. More specifically, we will differentiate between two cases: (i) the experimenter has no access to <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e046" xlink:type="simple"/></inline-formula> and only observed the decision maker's choices, <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e047" xlink:type="simple"/></inline-formula>, or (ii) the experimenter has partial knowledge of <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e048" xlink:type="simple"/></inline-formula> (to be defined more precisely later).</p>
<p>To illustrate our task setup further, consider a simple 2-AFC, in which the experimenter chooses at each trial the hidden state <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e049" xlink:type="simple"/></inline-formula> according to <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e050" xlink:type="simple"/></inline-formula> and <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e051" xlink:type="simple"/></inline-formula> (<xref ref-type="fig" rid="pone-0096511-g001">Fig. 1a</xref>). Based on this, one of two 3-sided coins (one fair, one biased) is chosen to generate the possible set of observations <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e052" xlink:type="simple"/></inline-formula>, either from coin 1 by <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e053" xlink:type="simple"/></inline-formula>, or from coin 2 by <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e054" xlink:type="simple"/></inline-formula>(see <xref ref-type="fig" rid="pone-0096511-g001">Fig. 1a</xref> for generative probabilities, parameterized by <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e055" xlink:type="simple"/></inline-formula>; <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e056" xlink:type="simple"/></inline-formula>). The decision maker observes the outcome of this coin flip, but does not know which coin was used to generate it. Assuming <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e057" xlink:type="simple"/></inline-formula> and <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e058" xlink:type="simple"/></inline-formula>, it is easy to show that the optimal strategy is to pick coin 1 (<inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e059" xlink:type="simple"/></inline-formula>) if <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e060" xlink:type="simple"/></inline-formula>, and coin 2 (<inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e061" xlink:type="simple"/></inline-formula>) otherwise (<xref ref-type="fig" rid="pone-0096511-g001">Fig. 1b</xref>, this corresponds to the maximum a-posterior estimate of the coin state; with <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e062" xlink:type="simple"/></inline-formula>, <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e063" xlink:type="simple"/></inline-formula> does not reveal anything about the hidden state, such that <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e064" xlink:type="simple"/></inline-formula> was chosen arbitrarily in this case). The experimenter, in contrast, only observes this decision, <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e065" xlink:type="simple"/></inline-formula>, but not the outcome <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e066" xlink:type="simple"/></inline-formula> of the coin flip. This abstract task contains all the essential ingredients of our framework and will be used throughout the text to illustrate important concepts.</p>
</sec><sec id="s2b">
<title>Relating Belief and Performance</title>
<p>To relate the belief of the decision maker to the performance observed by the experimenter, let us first define what exactly we mean by these measures. The ‘belief’ refers to the decision maker's belief at decision time of choosing the correct option <xref ref-type="bibr" rid="pone.0096511-Kepecs2">[20]</xref>. Thus, given observation <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e067" xlink:type="simple"/></inline-formula> and potential choice <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e068" xlink:type="simple"/></inline-formula>, this belief is the probability <disp-formula id="pone.0096511.e069"><graphic position="anchor" xlink:href="info:doi/10.1371/journal.pone.0096511.e069" xlink:type="simple"/><label>(1)</label></disp-formula></p>
<p>Here, we explicitly condition on the decision <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e070" xlink:type="simple"/></inline-formula> to make clear that we only consider observations <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e071" xlink:type="simple"/></inline-formula> that lead to decision <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e072" xlink:type="simple"/></inline-formula>. This conditioning is only hypothetical (“what is my belief if I were to choose <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e073" xlink:type="simple"/></inline-formula>”), such that the belief can be computed before a choice is performed. For the same reason, our analysis is easily generalized to the belief of un-chosen options, but to simplify exposition we restrict ourselves to the option that is finally chosen. In either case, the belief is a subjective probability, and available to the decision maker in every single trial.</p>
<p>The experimenter measures the decision maker's performance by the fraction of times that the correct choice was made. Thus, for a given hidden state <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e074" xlink:type="simple"/></inline-formula>, and assuming no knowledge of <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e075" xlink:type="simple"/></inline-formula>, the experimenter measures the probability that the decision maker chose <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e076" xlink:type="simple"/></inline-formula>, that is <disp-formula id="pone.0096511.e077"><graphic position="anchor" xlink:href="info:doi/10.1371/journal.pone.0096511.e077" xlink:type="simple"/><label>(2)</label></disp-formula>This performance measure is standard in the psychophysics and perceptual decision making literature <xref ref-type="bibr" rid="pone.0096511-Kiani1">[5]</xref>, <xref ref-type="bibr" rid="pone.0096511-Kepecs1">[8]</xref>, <xref ref-type="bibr" rid="pone.0096511-Wichmann1">[21]</xref>. It is a frequentist probability estimated by averaging over many trials in which <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e078" xlink:type="simple"/></inline-formula>, that is, trials in which the stimulus is maintained constant. This is, for instance, the measure that is plotted in psychometric curves for 2-alternative forced choice (2-AFC) task.</p>
<p>Given these definitions, we want to address how performance measured by the experimenter (Eq. (2)) relates to the decision maker's belief (Eq. (1)). As an intermediate step, we will first explore the condition under which performance equals belief <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e079" xlink:type="simple"/></inline-formula> averaged over observations <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e080" xlink:type="simple"/></inline-formula>, given by <disp-formula id="pone.0096511.e081"><graphic position="anchor" xlink:href="info:doi/10.1371/journal.pone.0096511.e081" xlink:type="simple"/><label>(3)</label></disp-formula>where the integral is over the full support of <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e082" xlink:type="simple"/></inline-formula>, that is, all possible values of <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e083" xlink:type="simple"/></inline-formula> that lead to choice <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e084" xlink:type="simple"/></inline-formula>. A joint probability decomposition of <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e085" xlink:type="simple"/></inline-formula> reveals that <disp-formula id="pone.0096511.e086"><graphic position="anchor" xlink:href="info:doi/10.1371/journal.pone.0096511.e086" xlink:type="simple"/><label>(4)</label></disp-formula>where <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e087" xlink:type="simple"/></inline-formula> and <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e088" xlink:type="simple"/></inline-formula> are the fractions of trials that the hidden state was <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e089" xlink:type="simple"/></inline-formula>, and <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e090" xlink:type="simple"/></inline-formula> was chosen, respectively. This equality shows that the performance is only equal to the average belief, that is <disp-formula id="pone.0096511.e091"><graphic position="anchor" xlink:href="info:doi/10.1371/journal.pone.0096511.e091" xlink:type="simple"/><label>(5)</label></disp-formula>if <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e092" xlink:type="simple"/></inline-formula>. In other words, Eq. (5) is only true when the frequency of choosing <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e093" xlink:type="simple"/></inline-formula> equals that of it being the correct choice. This is not always the case. For instance, these two probabilities differ in our 3-sided coin example (<xref ref-type="fig" rid="pone-0096511-g001">Fig. 1a</xref>), when choice <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e094" xlink:type="simple"/></inline-formula> is correct with probability <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e095" xlink:type="simple"/></inline-formula> and <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e096" xlink:type="simple"/></inline-formula>. In this case, if subjects pick the most likely choice, they will pick choice 1 with probability, <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e097" xlink:type="simple"/></inline-formula>. Clearly, <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e098" xlink:type="simple"/></inline-formula>, because choice 1 only occurs on 50% of the trials (<inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e099" xlink:type="simple"/></inline-formula>), but is picked by the subject over 83% (<inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e100" xlink:type="simple"/></inline-formula>) of the time. As a result, the decision maker's average belief will differ from the performance measures by the experimenter. In general, <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e101" xlink:type="simple"/></inline-formula> might hold for symmetric tasks with uniform priors over hidden states, but is likely to be violated in tasks that are asymmetric (for example, <xref ref-type="fig" rid="pone-0096511-g001">Fig. 1</xref>), or in which some choices are more likely to be correct on average than others.</p>
<p>To summarize, belief only equals performance when the frequency of choices matches the frequency of them being correct, and even then, this belief is the average belief across trials (Eq. (3)) in which a particular choice was made.</p>
</sec><sec id="s2c">
<title>Accumulation of evidence over time by diffusion/race models</title>
<p>Even though the established formalism is already able to capture simple experimental setups, its applicability is limited to cases where all the experimenter observes are the decision maker's choices, and nothing else (that is, the experimenter does not have access to <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e102" xlink:type="simple"/></inline-formula>). In general, the experimenter might have access to further information, such as the reaction time, that reveals additional details about the decision maker's state at decision time. Consider, for instance, a situation where the observation <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e103" xlink:type="simple"/></inline-formula> is a noisy version of an image drawn by the experimenter. In this case, clearly, the experimenter will have some, but only partial information about the decision maker's observation. A second important limitation of previous examples is that we have assumed the observation <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e104" xlink:type="simple"/></inline-formula> to be immediately available, whereas, usually, the decision maker needs to accumulate evidence over time before committing to a decision. In this and the next section we extend the previous formalism to fully accommodate in the theory these situations. In the following, we focus on diffusion and race models due to their popularity in cognitive sciences and neuroscience and their mathematical tractability. Despite this, we want to emphasize that our general theory on the relation of belief and performance remains valid even if the particular assumptions underlying these model choices (such as independent and identically distributed momentary evidence) are violated.</p>
<p>We start by considering a 2-AFC random dot reaction time task <xref ref-type="bibr" rid="pone.0096511-Britten1">[22]</xref>–<xref ref-type="bibr" rid="pone.0096511-Newsome1">[23]</xref>. At each trial, the experimenter chooses the motion direction (left or right) and coherence (fraction of dots moving coherently) which is subsequently used to generate the visual stimulus. The decision maker is told to identify as quickly and as accurately as possible the motion direction. In this task, the hidden state <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e105" xlink:type="simple"/></inline-formula> is the motion direction, while the coherence is a nuisance parameter that does not carry any information about the correct choice. The momentary evidence about <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e106" xlink:type="simple"/></inline-formula> in a short time window <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e107" xlink:type="simple"/></inline-formula> follows a Gaussian <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e108" xlink:type="simple"/></inline-formula> with mean <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e109" xlink:type="simple"/></inline-formula> and variance <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e110" xlink:type="simple"/></inline-formula>. Its mean rate <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e111" xlink:type="simple"/></inline-formula> is determined by the experimenter, and is positive for left-ward motion (<inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e112" xlink:type="simple"/></inline-formula>) and negative for right-ward motion (<inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e113" xlink:type="simple"/></inline-formula>), and its magnitude <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e114" xlink:type="simple"/></inline-formula> is proportional to the coherence of the random-dot motion. The decision maker can infer <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e115" xlink:type="simple"/></inline-formula> through the momentary evidence <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e116" xlink:type="simple"/></inline-formula>, which she can accumulate over time by a bounded drifting and diffusing particle <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e117" xlink:type="simple"/></inline-formula> with <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e118" xlink:type="simple"/></inline-formula>, where <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e119" xlink:type="simple"/></inline-formula> is a unit variance Gaussian white noise <xref ref-type="bibr" rid="pone.0096511-Ratcliff2">[24]</xref>–<xref ref-type="bibr" rid="pone.0096511-MorenoBote1">[27]</xref>. In this diffusion model (DM, <xref ref-type="fig" rid="pone-0096511-g002">Fig. 2a</xref>), <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e120" xlink:type="simple"/></inline-formula> is chosen if this particle hits the upper, potentially time-varying boundary at <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e121" xlink:type="simple"/></inline-formula>, that is <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e122" xlink:type="simple"/></inline-formula>, and <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e123" xlink:type="simple"/></inline-formula> is chosen if it hits the lower boundary at <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e124" xlink:type="simple"/></inline-formula>. We allow these boundaries to change with time to demonstrate the generality of our framework. Clearly, all principles discussed here transfer immediately to the more standard case of time-invariant boundaries. At the point when either of the boundaries has been reached, all the information required to compute the belief about the hidden state <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e125" xlink:type="simple"/></inline-formula> is the particle location at this time, that is <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e126" xlink:type="simple"/></inline-formula>, and the decision time <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e127" xlink:type="simple"/></inline-formula> (see Methods: 2-AFC decision making with diffusion models) <xref ref-type="bibr" rid="pone.0096511-Kiani1">[5]</xref>, <xref ref-type="bibr" rid="pone.0096511-Drugowitsch1">[26]</xref>. Thus, we define the observation <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e128" xlink:type="simple"/></inline-formula> as the pair particle location at decision and decision time, which are the sufficient statistics of this belief. In such a setup, the experimenter might be able to observe the time <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e129" xlink:type="simple"/></inline-formula> of this decision, but not necessarily the true state of the variable <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e130" xlink:type="simple"/></inline-formula>. This gives the experimenter partial knowledge of the state of the DM because knowing decision time <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e131" xlink:type="simple"/></inline-formula> tells the experimenter that one of the two bounds has been hit. More formally, knowing the decision time <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e132" xlink:type="simple"/></inline-formula>, the experimenter can restrict <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e133" xlink:type="simple"/></inline-formula> to the set <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e134" xlink:type="simple"/></inline-formula>, which denotes the set of observation vectors <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e135" xlink:type="simple"/></inline-formula> with decision time equal to <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e136" xlink:type="simple"/></inline-formula> which is simply the set in which the first component of the vector <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e137" xlink:type="simple"/></inline-formula> is either <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e138" xlink:type="simple"/></inline-formula> or <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e139" xlink:type="simple"/></inline-formula>. In fact, the experimenter can also infer whether the positive or the negative boundary was hit from observing the response of the subject, although the value of the boundary itself remains unknown. This partial knowledge can be exploited by the experimenter to get a better handle on the decision maker's belief, as we will describe further below.</p>
<fig id="pone-0096511-g002" position="float"><object-id pub-id-type="doi">10.1371/journal.pone.0096511.g002</object-id><label>Figure 2</label><caption>
<title>The diffusion model (DM) and 2-race model.</title>
<p>(a) In a DM, a particle drifts and diffuses over time. A decision is performed as soon as this particle reaches one of the two boundaries. The mean drift rate <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e140" xlink:type="simple"/></inline-formula>, which is unknown to the decision maker, determines which of the two choices is correct. In this illustration, the drift is towards the upper boundary, corresponding to hidden state <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e141" xlink:type="simple"/></inline-formula>, such that <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e142" xlink:type="simple"/></inline-formula> is the correct choice. We show eight (solid) trajectories leading to the correct choice (<inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e143" xlink:type="simple"/></inline-formula>) and two (dashed) trajectories leading to the wrong choice (<inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e144" xlink:type="simple"/></inline-formula>). Our framework allows for time-varying boundaries, as shown here and used to generate <xref ref-type="fig" rid="pone-0096511-g003">Figs. 3a/b</xref> and <xref ref-type="fig" rid="pone-0096511-g004">4a/b</xref>. (b) A race model features <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e145" xlink:type="simple"/></inline-formula> races (here <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e146" xlink:type="simple"/></inline-formula>) that compete against each other in a race towards a boundary of height <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e147" xlink:type="simple"/></inline-formula>. The race that first reaches its associated boundary determines the decision. The set of all races is described by a drifting/diffusing particle in <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e148" xlink:type="simple"/></inline-formula>-dimensional space. In our illustration this particle drifts towards the upper boundary (thus <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e149" xlink:type="simple"/></inline-formula>) and diffuses in both dimensions. Thus, four (solid) trajectories lead to the correct choice (<inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e150" xlink:type="simple"/></inline-formula>), and one (dashed) trajectory leads to the incorrect choice (<inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e151" xlink:type="simple"/></inline-formula>).</p>
</caption><graphic mimetype="image" xlink:href="info:doi/10.1371/journal.pone.0096511.g002" position="float" xlink:type="simple"/></fig>
<p>The same logic applies to scenarios in which more than two options are available to choose from. Let us consider a <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e152" xlink:type="simple"/></inline-formula>-AFC task for <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e153" xlink:type="simple"/></inline-formula> (<xref ref-type="fig" rid="pone-0096511-g002">Fig. 2b</xref> for <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e154" xlink:type="simple"/></inline-formula>). In this case, we assume that the experimenter presents a stimulus that determines <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e155" xlink:type="simple"/></inline-formula> non-negative drift rates <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e156" xlink:type="simple"/></inline-formula>. The hidden state is determined by the largest of these rates, such that <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e157" xlink:type="simple"/></inline-formula> if and only if all races <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e158" xlink:type="simple"/></inline-formula> feature a lower drift rate than race <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e159" xlink:type="simple"/></inline-formula>, that is, <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e160" xlink:type="simple"/></inline-formula>. The decision maker observes <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e161" xlink:type="simple"/></inline-formula> races, given by the drifting/diffusing particle <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e162" xlink:type="simple"/></inline-formula> starting at <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e163" xlink:type="simple"/></inline-formula>, towards a potentially time-varying boundary <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e164" xlink:type="simple"/></inline-formula> starting at <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e165" xlink:type="simple"/></inline-formula>. A decision strategy that maximizes the posterior belief under certain circumstances is to choose <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e166" xlink:type="simple"/></inline-formula> if race <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e167" xlink:type="simple"/></inline-formula> is the first to reach this boundary (see Methods: <italic>K</italic>-AFC decision making with race models). That is, <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e168" xlink:type="simple"/></inline-formula> if and only if <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e169" xlink:type="simple"/></inline-formula>, where <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e170" xlink:type="simple"/></inline-formula> is the first time at which either race has reach the boundary. Independent of the used decision strategy, it can be shown that the sufficient statistics that completely determine the decision maker's posterior belief about the hidden state are time <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e171" xlink:type="simple"/></inline-formula> and the particle locations <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e172" xlink:type="simple"/></inline-formula> at this time (see Methods: <italic>K</italic>-AFC decision making with race models) <xref ref-type="bibr" rid="pone.0096511-MorenoBote1">[27]</xref>. Thus, we define an observation in the race model setup to be these statistics at decision time <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e173" xlink:type="simple"/></inline-formula>, that is <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e174" xlink:type="simple"/></inline-formula>, where decision <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e175" xlink:type="simple"/></inline-formula> corresponds to <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e176" xlink:type="simple"/></inline-formula> and <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e177" xlink:type="simple"/></inline-formula> for all <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e178" xlink:type="simple"/></inline-formula>. The experimenter can again observes both the chosen option and the time of this choice, and so has partial access to the decision maker's observation <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e179" xlink:type="simple"/></inline-formula> by <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e180" xlink:type="simple"/></inline-formula>, where <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e181" xlink:type="simple"/></inline-formula> denotes all possible race states that result in a decision at time <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e182" xlink:type="simple"/></inline-formula> (which are all the vectors <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e183" xlink:type="simple"/></inline-formula> in which one of the first <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e184" xlink:type="simple"/></inline-formula> components is equal to <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e185" xlink:type="simple"/></inline-formula>). These examples illustrate that, despite our conceptually simple task formulation, we are able to capture a wide range of possible tasks and decision mechanisms that include non-uniform priors, and decisions that require the accumulation of evidence whose reliability might vary across trials.</p>
</sec><sec id="s2d">
<title>Relating belief and performance for partial knowledge of the observation</title>
<p>In the preceding cases, the experimenter has partial knowledge of the observation through observing the decision time. Here we describe how this information is used to refine the previously established relation between belief and performance. In general, we assume that partial knowledge of <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e186" xlink:type="simple"/></inline-formula> can be expressed by <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e187" xlink:type="simple"/></inline-formula>, which indicates that the experimenter knows that the observation has some features shared by all observations in <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e188" xlink:type="simple"/></inline-formula> (like, as the previous cases, the decision time), but does not know the observation <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e189" xlink:type="simple"/></inline-formula> itself. As a consequence, the performance as measured by the experimenter is given by <disp-formula id="pone.0096511.e190"><graphic position="anchor" xlink:href="info:doi/10.1371/journal.pone.0096511.e190" xlink:type="simple"/><label>(6)</label></disp-formula>where, when compared to Eq. (2), we additionally condition on <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e191" xlink:type="simple"/></inline-formula>. Hence, we assume that the experimenter evaluates the performance by binning trials by<inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e192" xlink:type="simple"/></inline-formula>. Setting <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e193" xlink:type="simple"/></inline-formula> (where <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e194" xlink:type="simple"/></inline-formula> is the set of all values that <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e195" xlink:type="simple"/></inline-formula> can take) recovers the original case in which the experimenter was unable to observe <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e196" xlink:type="simple"/></inline-formula>, demonstrating that the partial information case strictly generalizes the original case.</p>
<p>To relate belief and performance if partial knowledge is available, we again decompose the joint probability <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e197" xlink:type="simple"/></inline-formula> to get <disp-formula id="pone.0096511.e198"><graphic position="anchor" xlink:href="info:doi/10.1371/journal.pone.0096511.e198" xlink:type="simple"/><label>(7)</label></disp-formula>Thus, as before, performance only equals the average belief if <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e199" xlink:type="simple"/></inline-formula>, that is, if the fraction of choosing <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e200" xlink:type="simple"/></inline-formula> in trials in which <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e201" xlink:type="simple"/></inline-formula> equals the fraction of this choice being correct in such trials. Furthermore, the belief on the right-hand side of Eq. (7) is <disp-formula id="pone.0096511.e202"><graphic position="anchor" xlink:href="info:doi/10.1371/journal.pone.0096511.e202" xlink:type="simple"/><label>(8)</label></disp-formula>which is the trial-by-trial belief averaged over trials in which <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e203" xlink:type="simple"/></inline-formula> was chosen and <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e204" xlink:type="simple"/></inline-formula> holds. The integral is over the full support of <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e205" xlink:type="simple"/></inline-formula>, which is the subset of <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e206" xlink:type="simple"/></inline-formula> that leads to choice <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e207" xlink:type="simple"/></inline-formula>. Thus the same restrictions apply to the relation between belief and performance as when the experimenter does not know <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e208" xlink:type="simple"/></inline-formula>, only that now they relate to the subgroup of trials in which <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e209" xlink:type="simple"/></inline-formula>.</p>
</sec><sec id="s2e">
<title>Belief and Performance for Diffusion and Race Models</title>
<p>Returning to the example of the diffusion model, the decision maker's belief when choosing option 1 at time <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e210" xlink:type="simple"/></inline-formula> is <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e211" xlink:type="simple"/></inline-formula> (where observation <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e212" xlink:type="simple"/></inline-formula> is defined as <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e213" xlink:type="simple"/></inline-formula>) the performance measured by the experimenter is <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e214" xlink:type="simple"/></inline-formula>. Here <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e215" xlink:type="simple"/></inline-formula> denotes that the experimenter knows that a decision has been made at time <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e216" xlink:type="simple"/></inline-formula>, and <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e217" xlink:type="simple"/></inline-formula> implies – without specifying the height of the boundary – that option 1 has been chosen. We furthermore assume a symmetric prior on the drift rates, that is, <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e218" xlink:type="simple"/></inline-formula>. This implies for any decision time <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e219" xlink:type="simple"/></inline-formula> a uniform prior on hidden states, <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e220" xlink:type="simple"/></inline-formula>, and an equal probability of choosing either option, <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e221" xlink:type="simple"/></inline-formula>, such that the probability of choosing either option equals to it being correct, that is <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e222" xlink:type="simple"/></inline-formula>. Under these conditions we have previously established <xref ref-type="bibr" rid="pone.0096511-Drugowitsch1">[26]</xref> that performance equals average belief, such that <disp-formula id="pone.0096511.e223"><graphic position="anchor" xlink:href="info:doi/10.1371/journal.pone.0096511.e223" xlink:type="simple"/><label>(9)</label></disp-formula>Thus, the decision maker's belief when choosing option 1 at time <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e224" xlink:type="simple"/></inline-formula> equals her probability of making a correct choice at this time (<xref ref-type="fig" rid="pone-0096511-g003">Fig. 3a</xref>). It has not been shown before, however, that as soon as we start introducing asymmetry into the task by, for example, a non-uniform prior, this relationship will break down (<xref ref-type="fig" rid="pone-0096511-g003">Fig. 3b</xref>).</p>
<fig id="pone-0096511-g003" position="float"><object-id pub-id-type="doi">10.1371/journal.pone.0096511.g003</object-id><label>Figure 3</label><caption>
<title>Relationship between belief and performance in diffusion models (DMs) and race models.</title>
<p>(a) In a DM with uniform priors, <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e225" xlink:type="simple"/></inline-formula>, and symmetric boundaries, belief (data points) and performance (line) are equivalent. In the DM used to generate this figure, the boundaries collapse over time, causing a drop in belief/performance with time. If the boundaries were time-invariant instead, both belief and performance would be independent of time. (b) In the same DM with the same symmetric boundaries, but a non-uniform prior of <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e226" xlink:type="simple"/></inline-formula>, this equivalence fails to hold. It appears as if the decision maker were overconfident in her choices. (c) Simulations for a 2-race model with uniform priors, in which the winning race determines the choice, feature a strong fluctuation of the trial-by-trial belief around the decision maker's performance. It appears as if the decision maker features a belief that is idiosyncratic, fluctuating very strongly at each trial, although on average it equals her performance. In all panels the performance (with 95% CI) is estimated in bins of 250 ms, each containing data from 500 trials. The performance is measured as a fraction of trials in which option 1 was chosen when this choice was correct. For each of these bins, 10 examples (50 for the 2-race model) for the trial-by-trial belief when choosing option 1 are shown. This trial-by-trial belief is assumed to be either reported by the decision maker, or to be estimated from neural population activity. Details of how the models were simulated are in Methods: Generating <xref ref-type="fig" rid="pone-0096511-g003">Figures 3</xref> and <xref ref-type="fig" rid="pone-0096511-g004">4</xref>.</p>
</caption><graphic mimetype="image" xlink:href="info:doi/10.1371/journal.pone.0096511.g003" position="float" xlink:type="simple"/></fig>
<p>Interestingly, the belief averaged over all decisions made at time <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e227" xlink:type="simple"/></inline-formula> (Eq. (8)) in this example turns out to be equivalent to the belief held by the decision maker in each of these trials (Eq. (1)). Indeed, using our more general notation to express this, we have <disp-formula id="pone.0096511.e228"><graphic position="anchor" xlink:href="info:doi/10.1371/journal.pone.0096511.e228" xlink:type="simple"/><label>(10)</label></disp-formula>Thus, if the experimenter bins trials by decision time and computes the percentage of correct choices in each of these bins (as in <xref ref-type="fig" rid="pone-0096511-g003">Fig. 3a</xref>), this percentage will correlate perfectly with the decision maker's trial-by-trial belief at these decision times. In this model, the perfect correlation arises from to the lack of variability in decision confidence in this model, a result that will be violated in most general models (see below).</p>
<p>To understand why this property holds, it is instructive to revisit Eq. (8), which states that the average belief is the trial-by-trial belief held by the decision maker averaged over all trials in which choice <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e229" xlink:type="simple"/></inline-formula> was made, and <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e230" xlink:type="simple"/></inline-formula> specifies the time of this choice. For the diffusion model, knowing both choice and decision time corresponds to knowing which of the two boundaries was reached, and at which time, thus specifying the observation by <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e231" xlink:type="simple"/></inline-formula> and <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e232" xlink:type="simple"/></inline-formula> for <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e233" xlink:type="simple"/></inline-formula> and <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e234" xlink:type="simple"/></inline-formula>, respectively. Therefore, even if the bound height <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e235" xlink:type="simple"/></inline-formula> and thus the exact value of <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e236" xlink:type="simple"/></inline-formula> is unknown, the experimenter's knowledge of decision time and choice restricts <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e237" xlink:type="simple"/></inline-formula> to a single possible value, which results in the same belief every time this choice is made at this time. In general, as long as <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e238" xlink:type="simple"/></inline-formula> and <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e239" xlink:type="simple"/></inline-formula> restrict <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e240" xlink:type="simple"/></inline-formula> to a single possible observation, Eq. (10) holds. As a result, the diffusion model has the fortunate property that the experimenter has access to the trial-by-trial belief solely by measuring the performance of the decision maker. This has an important implication: for DMs applied to symmetric 2-AFC tasks, trial-by-trial belief, and not just averaged belief, equals performance, which is a very useful property for experimenters interested in inferring belief from performance <xref ref-type="bibr" rid="pone.0096511-Drugowitsch1">[26]</xref>.</p>
<p>This property is not shared by multiple-race models (<xref ref-type="fig" rid="pone-0096511-g003">Fig. 3c</xref>). In a multiple-race model as described above, the belief of the decision maker when choosing option 1 at time <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e241" xlink:type="simple"/></inline-formula> is her belief that the drift of the first race is larger than that of all other races, as given by <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e242" xlink:type="simple"/></inline-formula>, where we implicitly condition on no race having reached the boundary before <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e243" xlink:type="simple"/></inline-formula>. The performance as measured by the experimenter is the probability that option 1 was chosen at time <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e244" xlink:type="simple"/></inline-formula>, given that it was correct, as specified by <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e245" xlink:type="simple"/></inline-formula>, where <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e246" xlink:type="simple"/></inline-formula> implies that race 1 is the first to reach the boundary without specifying this boundary's height, and <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e247" xlink:type="simple"/></inline-formula>, where <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e248" xlink:type="simple"/></inline-formula>, denotes that some decision has been made at time <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e249" xlink:type="simple"/></inline-formula>. We furthermore assume that the prior <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e250" xlink:type="simple"/></inline-formula> has the same density for all permutations of the indices <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e251" xlink:type="simple"/></inline-formula> on the <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e252" xlink:type="simple"/></inline-formula>'s, such that <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e253" xlink:type="simple"/></inline-formula> for all <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e254" xlink:type="simple"/></inline-formula>. Under these conditions, we can again relate performance and average belief by <disp-formula id="pone.0096511.e255"><graphic position="anchor" xlink:href="info:doi/10.1371/journal.pone.0096511.e255" xlink:type="simple"/><label>(11)</label></disp-formula>However, in contrast to the DM, the average belief, on the right-hand side of Eq. (11) is not equal to the trial-by-trial belief as held by the decision maker. This discrepancy stems from the decision maker's belief not only depending on the state of the winning race, but also on that of all other races. For example, all races being close the boundary would induce higher uncertainty about the correctness of the decision than if there is a clear separation between the winning and the losing races (see also Eq. (25)). As a result, this belief varies across trials even if the same decision is made at the same time. Thus, the experimenter is unable to determine the decision maker's trial-by-trial belief by measuring her performance, but only its average. More formally, the probability <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e256" xlink:type="simple"/></inline-formula> that specifies in Eq. (8) which trials the belief is averaged over, now has non-zero probability for multiple values of <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e257" xlink:type="simple"/></inline-formula>. This is because <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e258" xlink:type="simple"/></inline-formula> and <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e259" xlink:type="simple"/></inline-formula> specify the winning race and bound-hitting time respectively, but the state of the losing races are only restricted to be somewhere below the decision threshold. Thus, these can take any state as long as <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e260" xlink:type="simple"/></inline-formula> and <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e261" xlink:type="simple"/></inline-formula> hold. As a result, the average is computed over all possible states of the losing races that satisfy <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e262" xlink:type="simple"/></inline-formula> and <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e263" xlink:type="simple"/></inline-formula>, causing the average belief to differ from the decision maker's trial-by-trial belief. As we will show later, this is a general property of all decision making procedures in which the decision maker's belief depends on decision variables that are not accessible to the experimenter.</p>
<p>In the example in <xref ref-type="fig" rid="pone-0096511-g003">Fig. 3c</xref>, the Pearson correlation coefficient between the binned percentage of corrected trials and the decision maker's trial-by-trial belief drops from close to one for the diffusion model to around 0.18 for the 2-race model. With less than 200 trials worth of observations, such a correlation coefficient is not even considered significantly different from zero at the 0.01 level. This illustrates that, in practice, such fluctuations can seriously impair the relation between trial-by-trial belief and actual performance.</p>
</sec><sec id="s2f">
<title>Relevance for Decision Maker</title>
<p>We have established that the decision maker's performance equals her belief only in rare cases, even if we assume that the decision maker holds the correct model of the environment. For instance, if the probability of the choices is not uniform, or subjects shows biases or preferences for a particular choice, belief and performance are not expected to coincide. The equality between belief and performance depends not only on the decision maker's strategy to perform the decision (that is, the used decision model, e.g. with biases or not), but also on the task that the decision maker has to solve (e.g. with or without non-uniform priors on the correct choices). The dissociation between belief and performance in most natural conditions therefore seems to violate the very assumption that the subjects have a correct model of the world since her own belief does not predict performance.</p>
<p>Yet, let us reconsider the quantity that the decision maker should monitor to feature efficient behavior. A belief (e.g. 0.8) is a useful quantity only to the extent that it predicts the percentage of time (e.g. 80%) the subject will be correct every time she observes <italic>x</italic> and decision <italic>k</italic> was taken, which is simply the quantity <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e264" xlink:type="simple"/></inline-formula>. This is the same quantity we have defined as the ‘belief’ of the subject in <xref ref-type="disp-formula" rid="pone.0096511.e069">equation (1</xref>). To compute this quantity, the subject needs to use Bayes rule, which relies on knowledge of the true generative model <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e265" xlink:type="simple"/></inline-formula> and prior <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e266" xlink:type="simple"/></inline-formula>. When this is the case, the belief computed by the subject will be exactly equal to <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e267" xlink:type="simple"/></inline-formula>, that is, equal to the percentage of time she will be correct whenever she observes <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e268" xlink:type="simple"/></inline-formula> and made decision <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e269" xlink:type="simple"/></inline-formula>. Therefore, although we have gained crucial insights into the decision process with the study of the relationship between performance and belief, the quantity we have called performance, <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e270" xlink:type="simple"/></inline-formula>, which is commonly measured by experimentalists, is not directly relevant to the decision maker's self-monitoring of her efficiency.</p>
<p>We can gain further insight into the sufficiency of monitoring ones belief by reconsidering the relationship we use to establish the equivalence between belief and performance. If we sum both sides of Eq. (7) over all <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e271" xlink:type="simple"/></inline-formula>, we trivially find <disp-formula id="pone.0096511.e272"><graphic position="anchor" xlink:href="info:doi/10.1371/journal.pone.0096511.e272" xlink:type="simple"/><label>(12)</label></disp-formula>showing that the average belief over all choices on the left-hand side equals the average performance over all hidden states on the right-hand side, even when <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e273" xlink:type="simple"/></inline-formula>, that is, even if the decision-maker does not perform frequency matching. Thus, as soon as we stop conditioning on choice or hidden state, we regain equality under all conditions. The inequality due to conditioning arose from considering a different set of trials for belief than for performance by conditioning on information unavailable to the decision maker (that is, the hidden state). Regaining equality once we consider the same set of trials confirms that monitoring ones belief will indeed provide a correct picture of ones behavioral efficiency, but only on average.</p>
<p>Note however that even when belief and performance are not equivalent, they are positively and linearly related <italic>on average</italic>. To see this, observe that in Eq. (4) both the choice probability <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e274" xlink:type="simple"/></inline-formula> and the prior probability <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e275" xlink:type="simple"/></inline-formula> are constant across trials, such that an increase in the average belief <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e276" xlink:type="simple"/></inline-formula> directly relates to an increase in performance <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e277" xlink:type="simple"/></inline-formula>. This also holds for the more general case in which we condition on a subset of observations, as in Eq. (7). As a result, the decision maker can use the average belief's gradient to improve her performance even in cases where these two quantities are not equivalent. Still, one again should be aware that this linear relationship holds only on average, such that – depending on how strongly the trial-by-trial belief fluctuates around the average belief, as shown above – this relationship might be of limited use.</p>
</sec><sec id="s2g">
<title>Relevance to Experimenter</title>
<p>From the experimenter's perspective, an equality between belief and performance is important as it would imply that one could use performance as a surrogate for belief (or average belief). Thus, experimenters might be tempted to avoid more complex experimental setups in which these two quantities are not equal, since it would become unclear how to assess the decision maker's belief. Yet, a simple remedy presents itself by considering what needs to be known to evaluate average belief directly. Average belief, <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e278" xlink:type="simple"/></inline-formula>, is the probability that the hidden state was <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e279" xlink:type="simple"/></inline-formula> when subject chose <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e280" xlink:type="simple"/></inline-formula> and <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e281" xlink:type="simple"/></inline-formula>. From a frequentist point of view, this is the percentage of time the subject made the correct choice (that is subject chose <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e282" xlink:type="simple"/></inline-formula> when the hidden state was indeed <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e283" xlink:type="simple"/></inline-formula>) given partial knowledge<inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e284" xlink:type="simple"/></inline-formula>. Therefore, if we bin all the trials for which the subject chose k and <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e285" xlink:type="simple"/></inline-formula>, the percentage of correct responses will converge to <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e286" xlink:type="simple"/></inline-formula> for very large number of trials. More formally: <disp-formula id="pone.0096511.e287"><graphic position="anchor" xlink:href="info:doi/10.1371/journal.pone.0096511.e287" xlink:type="simple"/><label>(13)</label></disp-formula>where the sums are over all trials, indexed by <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e288" xlink:type="simple"/></inline-formula>, and <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e289" xlink:type="simple"/></inline-formula> is the identifier function that returns <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e290" xlink:type="simple"/></inline-formula> is the statement <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e291" xlink:type="simple"/></inline-formula> is true, and <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e292" xlink:type="simple"/></inline-formula> otherwise. This shows that the experimenter can evaluate the decision maker's average belief, even when belief and performance do not correspond to each other, as illustrated in <xref ref-type="fig" rid="pone-0096511-g004">Fig 4a</xref>. However, even then, this average belief might only be weakly correlated with the decision maker's trial-by-trial belief (for example, <xref ref-type="fig" rid="pone-0096511-g004">Fig. 4b</xref>), such that this average belief might tell the experimenter little about the decision maker's belief in individual trials.</p>
<fig id="pone-0096511-g004" position="float"><object-id pub-id-type="doi">10.1371/journal.pone.0096511.g004</object-id><label>Figure 4</label><caption>
<title>Comparing estimated belief with performance and trial-by-trial belief.</title>
<p>(a) A DM with a non-uniform prior of <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e293" xlink:type="simple"/></inline-formula> as in <xref ref-type="fig" rid="pone-0096511-g003">Fig.  3b</xref>. Trial-by-trial belief differs from performance because of the asymmetric prior. By contrast, the estimated belief using Eq. (13) matches the trial-by-trial belief, because the decision maker's state is fully observable in a DM. (b) A two race model with uniform priors as in <xref ref-type="fig" rid="pone-0096511-g003">Fig 3c</xref>. This time, the decision maker's state is not fully observable because the state of the losing race is unknown to the experimenter. As a consequence, the belief estimated by Eq. (13) no longer matches the trial-by-trial belief of the observer but only the averaged belief, where the average is performed over the state of the losing race. Details of the model simulations are described in Methods: Generating <xref ref-type="fig" rid="pone-0096511-g003">Figures 3</xref> and <xref ref-type="fig" rid="pone-0096511-g004">4</xref>.</p>
</caption><graphic mimetype="image" xlink:href="info:doi/10.1371/journal.pone.0096511.g004" position="float" xlink:type="simple"/></fig>
<p>To summarize, the relevant quantity for estimating belief is not performance as defined by the psychometric curves, but the percentage of correct responses conditioned on the subject response and partial knowledge of <italic>x</italic> (for example, percentage of correct response given that the subject chose rightward motion and the reaction time is between <italic>t</italic> and <italic>t</italic>+<italic>dt</italic>). In a psychometric curve, the percentage correct is conditioned on the true state of the world (for example, actual motion was to the right), while we are now conditioning on the decision maker's response. Note that this is the same fix as the one we used in the previous section when we considered the point of view of the decision maker.</p>
</sec><sec id="s2h">
<title>The hard-easy effect in psychometric curves</title>
<p>In general, the relation between belief and performance breaks down as soon as performance is measured conditional on events that are fundamentally inaccessible either to the experimenter or the decision maker, that is, in the case of information asymmetry. This breakdown could explain a conspicuous result known as the <italic>hard-easy</italic> effect: when asked to estimate their confidence in a judgment, subjects tend to overestimate their confidence on hard trials and to underestimate their confidence on easy trials <xref ref-type="bibr" rid="pone.0096511-Lichtenstein1">[17]</xref>, <xref ref-type="bibr" rid="pone.0096511-Erev1">[28]</xref>–<xref ref-type="bibr" rid="pone.0096511-Pfeifer1">[29]</xref>. To see how such an effect could arise from this breakdown, let us consider a simple reaction time task, for example the random dot motion task described before, whose difficulty varies between trials. We represent this difficulty by, at the beginning of each trial, drawing <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e294" xlink:type="simple"/></inline-formula> from a point-wise distribution shown in <xref ref-type="fig" rid="pone-0096511-g005">Fig. 5a</xref>, corresponding to a task in which the difficulty is interleaved across trials and can take one of a fixed number of alternatives. Here, the sign of <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e295" xlink:type="simple"/></inline-formula> determines the hidden state <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e296" xlink:type="simple"/></inline-formula>, and <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e297" xlink:type="simple"/></inline-formula> specifies the trial's difficulty (that is, the dot motion's coherence), with smaller <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e298" xlink:type="simple"/></inline-formula>'s corresponding to harder trials <xref ref-type="bibr" rid="pone.0096511-Drugowitsch1">[26]</xref>. The range of possible <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e299" xlink:type="simple"/></inline-formula>'s controls the average difficulty of the task. A standard practice in such setups is to bin trials by their difficulty <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e300" xlink:type="simple"/></inline-formula> and plot the average reaction time and fraction of correct choices for each of these bins separately (the so-called chronometric and psychometric curves, respectively). Using standard analytical results for the first-passage time and choice probability for diffusion models in which <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e301" xlink:type="simple"/></inline-formula> determines the drift rate (see Methods: Computing belief in a drift diffusion model with varying difficulty) leads to the chronometric and psychometric curve shown in <xref ref-type="fig" rid="pone-0096511-g005">Fig. 5b</xref>. Here, we have chosen a diffusion model with time-invariant boundaries, as the assumption of a trial-by-trial change in task difficulty causes the belief at the boundary to be time-dependent even when the boundary is not. Our conclusions do not depend on this choice, as the same principles apply to the case of time-dependent boundaries.</p>
<fig id="pone-0096511-g005" position="float"><object-id pub-id-type="doi">10.1371/journal.pone.0096511.g005</object-id><label>Figure 5</label><caption>
<title>Mismatch between average belief and performance when conditioning on task difficulty: the hard-easy effect and miscalibration.</title>
<p>We simulated a task with varying difficulty given by a diffusion model with a drift rate whose magnitude and sign varied across trials, while being constant within each trial. (a) The top graph shows the across-trials point-wise prior on the drift rate used in the simulation that roughly approximates a zero-mean Gaussian (dashed line). We computed the decision maker's belief by either using this point-wise prior directly, or by assuming it to follow a too-wide zero-mean Gaussian (dotted line). The bottom graph shows that the point-wise prior corresponds to the 10<sup>th</sup>, 20<sup>th</sup>, …, 90<sup>th</sup> percentile of the Gaussian it approximates. (b) The decision maker's chronometric (top) and psychometric (bottom) function over task difficulty (magnitude of <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e302" xlink:type="simple"/></inline-formula>) for non-negative drift rates. Correct choices here correspond to hitting the upper bound of the diffusion model if the drift rate is positive, and the lower bound otherwise. The bottom graph also shows the decision maker's average belief over <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e303" xlink:type="simple"/></inline-formula> for both correct and error trials (dots exactly one top of each other, as confidence for correct and error trials is identical) based on the correct, point-wise prior (squares, +/− 2SD) and on the incorrect Gaussian prior (crosses). In both cases, the mismatch between average belief and performance when conditioning on task difficulty is clearly visible. (c) The calibration curves, showing the probability of performing correct choices as a function of the decision maker's belief. When binning trials by difficulty (that is, drift rate magnitude), this choice probability is constant while the decision maker's belief varies across trials. This results in flat calibration curves (dashed/dotted lines), caricaturizing the frequently observed hard-easy effect. Once we stop conditioning on task difficulty, the calibration curve reveals perfect calibration (solid line). (d) Calibration curves for a mismatch between the actual distribution of task difficulties and that assumed by the decision maker to compute her belief. We consider the case in which the decision maker's distribution is too narrow (that is, has too small standard deviation; dotted line) or too wide (too large standard deviation; solid line). Both cases feature a clear miscalibration of the decision maker's belief.</p>
</caption><graphic mimetype="image" xlink:href="info:doi/10.1371/journal.pone.0096511.g005" position="float" xlink:type="simple"/></fig>
<p>Intuitively, one would expect the fraction of correct choices, as shown by psychometric curve, to be a good predictor of the decision maker's belief. However, comparing it to the across-trial average of the optimally computed belief (Eq. (28), shown in <xref ref-type="fig" rid="pone-0096511-g005">Fig. 5b</xref>) reveals this to be a fallacy. More specifically, the performance varies widely as a function of difficulty, while the average belief is only very weakly related to this difficulty. This is confirmed by a correlation coefficient below 0.35 between the psychometric curve and the trial-by-trial belief.</p>
<p>As before, the origin of the difference between belief and performance lies in conditioning the performance measure on an event that is fundamentally inaccessible to the decision maker, in this case the trial-by-trial difficulty <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e304" xlink:type="simple"/></inline-formula> (although this time we are assuming that the experimenter knows more than the subject, as opposed to the converse). In this experiment, the decision maker does not know this difficulty, which is varied from trial to trial, and so needs to rely on the prior distribution (<xref ref-type="fig" rid="pone-0096511-g005">Fig. 5a</xref>) across trials to infer her belief. This leads to overconfidence in hard trials, and underconfidence in easy trials (left-most and right-most point in <xref ref-type="fig" rid="pone-0096511-g005">Fig. 5b</xref>, respectively). Consider, for example, trials in which <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e305" xlink:type="simple"/></inline-formula> (corresponding to 0% coherence in the random dot task), such that performance is, by definition, at chance. Nonetheless, random fluctuations in the stimulus cause the decision maker to decide for one of the two options, at which point her belief about the decision's correctness will be above chance. In fact, it can be shown that a belief of 0.5 will only ever occur for the impossible case of infinite decision times (Eq (28)). As a consequence, the decision maker's belief for trials in which <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e306" xlink:type="simple"/></inline-formula> will be above her average performance in these trials, which, from the experimenter's point-of-view, leads to overconfidence. A similar argument explains the underconfidence for trial difficulties in which the decision maker features close-to-perfect performance. Thus, even though by Eq. (12) the belief equals performance when averaged across all difficulties, assessing this equality while conditioning on trial difficulty makes this equality seem violated. This last point is particularly important in the light of claims that this <italic>hard-easy effect</italic> might be grossly over-estimated due to simply being an artifact of binning or measuring performance by averaging over binary choices <xref ref-type="bibr" rid="pone.0096511-Juslin1">[16]</xref>, <xref ref-type="bibr" rid="pone.0096511-Merkle1">[30]</xref>. In our case, it instead stems from conditioning the decision-makers reported belief and observed performance on variables that are not readily available to the subject. Although we have shown this result for a particular example of a diffusion model with time-independent decision bounds, our results are generally valid also for diffusion models with time-dependent bounds and race models. As we shown next, this effect could also arise even when performance is not conditioned on task difficulty, but the subjects assume the wrong prior over task difficulty.</p>
</sec><sec id="s2i">
<title>Miscalibration due to the mismatch between experimenter's and decision-maker's prior: signatures of suboptimal priors</title>
<p>Calibration of confidence judgments is usually assessed by the calibration curve <xref ref-type="bibr" rid="pone.0096511-Moore1">[18]</xref>, <xref ref-type="bibr" rid="pone.0096511-Baranski1">[31]</xref>–<xref ref-type="bibr" rid="pone.0096511-Ferrell2">[33]</xref>, which results from binning trials by the reported confidence and then plotting the fraction of correct trials for each bin. For perfectly calibrated decision makers, the fraction of correct trials ought to correspond to their confidence, in which case the calibration curve follows the identity line. If we perform the same analysis on the simulated behavior conditional on task difficulty in the example described in the previous section, we find strong deviations from this identity line that reflect the corresponding over- and underconfidence for easy and hard trials, respectively (<xref ref-type="fig" rid="pone-0096511-g005">Fig 5c</xref>, dashed/dotted lines; compare belief with performance in <xref ref-type="fig" rid="pone-0096511-g005">Fig 5b</xref>, bottom). In contrast, if we cease to condition on difficulty and analyze the whole dataset at once, we find perfect calibration (<xref ref-type="fig" rid="pone-0096511-g005">Fig. 5c</xref>, solid line), as predicted by Eq. (12). This again demonstrates that, as long as the belief is computed from the correct generative model (that is, in a Bayes-optimal way), average belief will equal average performance.</p>
<p>If a Bayes-optimal model of decision making produces perfect calibration, it follows that a calibration mismatch implies that subjects deviates from Bayes optimality. There are several methods available for detecting such deviations. For instance, in the <italic>decision variable partition model</italic> <xref ref-type="bibr" rid="pone.0096511-Ferrell1">[32]</xref>–<xref ref-type="bibr" rid="pone.0096511-Ferrell2">[33]</xref>. the experimental data are used to estimate the function employed by the decision maker to map internal observations, <bold><italic>x</italic></bold>, onto belief. This function can then be compared to the Bayes optimal function to determine whether subjects are miscalibrated (see Methods: Modeling miscalibrations by the decision variable partition model). The problem with this approach is that it does not provide an explanation for why subject use a suboptimal function, a problem shared by other models <xref ref-type="bibr" rid="pone.0096511-Vickers1">[7]</xref>, <xref ref-type="bibr" rid="pone.0096511-Merkle2">[34]</xref>.</p>
<p>One possibility is that subjects do not know the generative model perfectly. For example, subjects would be miscalibrated if they use the wrong prior over task difficulty. This is a very likely situation as subjects have to learn the distribution of trial difficulties used by the experimenter, a process that would take much longer than the duration of the experiments. This effect is illustrated in <xref ref-type="fig" rid="pone-0096511-g005">Fig. 5d</xref> which compares the calibration curves for a model using the true prior over task difficult and one assuming a much wider (or much narrower) distribution than the true one. In this case, the model exhibits clear deviation from perfect calibration. Therefore, miscalibration could be due in part to imperfect knowledge of the generative model. This potential explanation for miscalibration has already been suggested conceptually in <xref ref-type="bibr" rid="pone.0096511-Gigerenzer1">[12]</xref>, but here we made its statement more quantitative.</p>
</sec><sec id="s2j">
<title>Average versus trial-by-trial belief</title>
<p>One important caveat to the experimenter's access to the decision maker's belief, as for example by utilizing Eq. (13), is that this belief can only be measured on average rather than trial-by-trial. This is a result of the experimenter's inability to observe <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e307" xlink:type="simple"/></inline-formula> in general, causing an asymmetry in the information held by decision maker and experimenter. As shown before, the use of DMs in 2-AFC tasks do not cause such an asymmetry, as at decision time it is known that the diffusing particle has reached the boundary. In race models, in contrast, the state of the losing races is unknown, such that the belief computed with Eq. (13) does not correspond to the trial-by-trial belief (<xref ref-type="fig" rid="pone-0096511-g004">Fig. 4b</xref>) but only to the belief averaged over the unobserved state of the loosing races. As already pointed out above, this causes the trial-by-trial belief to be only weakly correlated with average performance – a correlation that might even be missed if the number of observed trials is low.</p>
<p>The same issues come up when considering the Sequential Probability Ratio Test (SPRT) <xref ref-type="bibr" rid="pone.0096511-Wald1">[35]</xref>–<xref ref-type="bibr" rid="pone.0096511-Wald2">[36]</xref> and its multi-hypothesis (that is, <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e308" xlink:type="simple"/></inline-formula>) variants (MSPRTs) <xref ref-type="bibr" rid="pone.0096511-Fishman1">[37]</xref>–<xref ref-type="bibr" rid="pone.0096511-Dragalin1">[40]</xref>. The SPRT, which has been shown to yield the optimal speed/accuracy trade-off for 2-AFC tasks with a single known task difficulty <xref ref-type="bibr" rid="pone.0096511-Wald2">[36]</xref>, is based on accumulating the relative evidence for one option over the other up to a time-invariant boundary, at which a decision is made. This boundary specifies the belief at decision time, such that the same belief is held every time a decision is made. In other words, the average belief at the boundary is equivalent to the trial-by-trial belief, similar to the DM. The MSPRTs, on the other hand, only feature an optimal speed/accuracy trade-off in some asymptotic sense. They exist in several variants that are all based on continuously updating the posterior belief of all options but differ in how they specify the decision bounds. Variants that commit to a decision as soon as the highest posterior belief across options has reached a pre-set threshold <xref ref-type="bibr" rid="pone.0096511-Fishman1">[37]</xref>–<xref ref-type="bibr" rid="pone.0096511-Baum1">[38]</xref>, <xref ref-type="bibr" rid="pone.0096511-Dragalin1">[40]</xref> will feature the same belief across all trials, just as the DM. In contrast, if their decision threshold becomes a function of the beliefs for various options <xref ref-type="bibr" rid="pone.0096511-Lorden1">[39]</xref>–<xref ref-type="bibr" rid="pone.0096511-Dragalin1">[40]</xref>, their belief in the correctness of the chosen option might vary across trials, as in race models.</p>
<p>In general, the trial-by-trial belief differs from the average belief as soon as the minimal sufficient statistics of the decision maker's belief fluctuate at decision time, even if the experimenter bins trials according to all available information, such as choice and decision time (for a more formal statement see Methods: Equivalence of average and trial-by-trial belief). For DMs, the sufficient statistics are fully determined by the aforementioned measures, but for race models these measures are not sufficiently restrictive. It might seem that this is due to the larger number of possible choices for the race model. However, it is erroneous to attribute the difference between DMs and race models solely to the number of choices. Consider, for example, an orientation categorization task in which the observed orientation is a noisy instantiation of the orientation associated with one of the <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e309" xlink:type="simple"/></inline-formula> generative categories. In this case, the minimal sufficient statistics is the perceived orientation, which can be represented by a scalar value. Even if we increase the number of possible categories and with it the number of possible options to choose from, the dimensionality of the minimal sufficient statistics remains unchanged (see Methods: Minimal sufficient statistics in an orientation categorization task). Rather, what matters is the number of independent sources that ambiguously generate the observations. While in diffusion models, only a single such source exists, a race model with <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e310" xlink:type="simple"/></inline-formula> races assumes <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e311" xlink:type="simple"/></inline-formula> such sources. In the categorization task, in contrast, the sole source of information is the observed orientation, which does not depend on the number of possible choices. Thus, if the experimenter aims at estimating the decision maker's trial-by-trial belief, it is important to design experiments that control and restrict number and nature of these sources.</p>
</sec><sec id="s2k">
<title>Generalizations</title>
<p>Our findings are robust to changes in the details of the framework. One could, for example, imagine that the decision is stochastically rather than deterministically based on <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e312" xlink:type="simple"/></inline-formula> through <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e313" xlink:type="simple"/></inline-formula>. Furthermore, we could assume that the experimenter has partial knowledge of <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e314" xlink:type="simple"/></inline-formula> through a two-step generative model, <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e315" xlink:type="simple"/></inline-formula> (for example, a generated image) and <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e316" xlink:type="simple"/></inline-formula> (for example, the neural response to that image), where the experimenter observes <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e317" xlink:type="simple"/></inline-formula> and the decision maker makes a decision based on <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e318" xlink:type="simple"/></inline-formula>. While either of these modifications changes the details of the formulation, belief still only corresponds to performance if task and prior are symmetric, and is in most cases only measurable by the experimenter on average.</p>
<p>Extending the framework to value-based decision making might be possible, and is mandatory for a complete theory of belief and its relation to choice. However, assigning different values to different choices introduces ambiguity about which decisions ought to be considered correct and which are incorrect. Thus, several definitions of belief and performance might be possible. For this reason, we restricted our exposition to the case in which a clear definition of “correct” and “incorrect” exists.</p>
</sec></sec><sec id="s3">
<title>Discussion</title>
<p>We have described how the performance of a decision maker (defined as the fraction of correct responses given the world's true state) relates to its belief of having made the correct decision, and the relevance of this relation for both the decision maker's self-monitoring and an experimenter interested in the decision maker's belief. Specifically, we have shown that performance only equals belief in cases where these measures are conditioned on quantities that are known to both the experimenter and the decision maker. This equality starts breaking down in case of information asymmetry between decision maker and experimenter. One such asymmetry occurs if the experimenter conditions performance on the true state of the world, which is unknown to the decision maker. In this case performance only equals belief for symmetric tasks, in which the probability of choosing a particular option equals the probability of this choice to be correct. Even then, the equality only holds for the average belief across many trials, while the decision maker's belief per trial might fluctuate around this average. This is the result of another information asymmetry, in which the experimenter is unable to access the decision maker's internal state at decision time, and so has to average over it. Furthermore, we have discussed that the decision maker can evaluate how well she performs the task even if her belief does not equal her performance. This is because the relevant quantity for self-monitoring is belief, computed as the expectation that the decision maker was correct given her response, rather than performance, computed as the fraction of times a decision maker was correct given the state of the world predetermined by the experimenter. Also, the experimenter does not need to measure performance to assess the decision maker's belief, as the latter is directly measurable at least on average as the fraction of times that the decision maker was correct given her choice, assuming that the decision maker has the correct model of the task. Similarly to the relation between belief and performance, however, this belief can in most cases only be computed on average, around which the decision maker's trial-by-trial belief fluctuates.</p>
<p>To relate belief and performance, we have assumed the decision maker to have fully learned the generative model of the task. In other words, the decision maker is able to infer optimally the posterior distribution over each of the choices being correct. While this might be a valid assumption in well-trained, low-level tasks, such as detecting a flash of light in an otherwise dark room, it is most certainly violated in more complex, high-level, decision making <xref ref-type="bibr" rid="pone.0096511-Beck2">[41]</xref>–<xref ref-type="bibr" rid="pone.0096511-Kahneman1">[42]</xref>. As we have seen, partial learning of the generative model of the task could lead to a mismatch between belief and performance, and could explain in particular the hard-easy effect (i.e. overconfidence for near-chance performance, underconfidence for high performance). This effect might arise in particular from assuming that the prior distribution over task difficulty is wider than it really is.</p>
<p>We have also seen that, even for rational decision-makers with a perfect knowledge of the task, the hard-easy effect arises naturally if the experimenter conditions performance and belief on trial difficulty when plotting the psychometric curve: as shown in <xref ref-type="fig" rid="pone-0096511-g005">Fig. 5b</xref>, rational decision-makers will seem underconfident in easy trials, and overconfident in difficult trials. We have identified this mismatch to result from the experimenter conditioning on variables of the task (as trial difficulty <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e319" xlink:type="simple"/></inline-formula> in diffusion models) that are fundamentally inaccessible to the decision maker, who instead can only rely on her prior over trial difficulties. Thus, the mismatch emerges again due to an information asymmetry between decision maker and experimenter.</p>
<p>Therefore, the hard-easy effect could be due to either subjects using the wrong generative model, or the experimenter assuming more knowledge than is available to the subject. Our proposal differs from a related one in <xref ref-type="bibr" rid="pone.0096511-Moore1">[18]</xref> where the hard-easy effect is explained by subjects assuming a single, certain, but biased task difficulty. We, in contrast, assume that the subject's uncertainty about this task difficulty is to blame.</p>
<p>For all of the above we want to emphasize that most of the literature on the calibration of confidence judgments is based on explicit, e.g. verbal, reporting of this confidence <xref ref-type="bibr" rid="pone.0096511-Lichtenstein1">[17]</xref>–<xref ref-type="bibr" rid="pone.0096511-Moore1">[18]</xref>, <xref ref-type="bibr" rid="pone.0096511-Baranski1">[31]</xref>, <xref ref-type="bibr" rid="pone.0096511-Zylberberg1">[43]</xref> which could also contribute to miscalibration of confidence. There is indeed clear evidence for the existence and use of uncertainty information about task-relevant variables in multisensory information <xref ref-type="bibr" rid="pone.0096511-Ernst1">[3]</xref>, <xref ref-type="bibr" rid="pone.0096511-Battaglia1">[44]</xref>, post-decision wagering <xref ref-type="bibr" rid="pone.0096511-Kiani1">[5]</xref>, and related paradigms <xref ref-type="bibr" rid="pone.0096511-Kepecs2">[20]</xref>. However, it is less clear if this information is accurately accessible for explicit reporting, or if this reporting is not part of the normal decision-making repertoire, but instead needs to be learned as a separate task, thus justifying models with a confidence judgment process that is at least to some degree separate from that leading to decisions <xref ref-type="bibr" rid="pone.0096511-Pleskac1">[15]</xref>, <xref ref-type="bibr" rid="pone.0096511-Galvin1">[45]</xref>–<xref ref-type="bibr" rid="pone.0096511-Maniscalco1">[46]</xref>. Either option might introduce additional biases <xref ref-type="bibr" rid="pone.0096511-Juslin1">[16]</xref>, <xref ref-type="bibr" rid="pone.0096511-Merkle1">[30]</xref>, such that it remains to be seen if the observed deviations from perfect is a property of the underlying inference process leading to the decision maker's belief, as we have suggested, or simply a property of the mapping of confidence onto explicit reports. In light of this, it seems advisable to assess this belief more directly by behavioral measures rather than by explicit reports.</p>
<p>Having identified some of the possible fallacies that can occur when relating belief and performance, we can revisit previously mentioned illustrative work on the decision maker's belief. In <xref ref-type="bibr" rid="pone.0096511-Kiani1">[5]</xref>, for example, it at first appears as if the authors wrongly condition on the task difficulty (in their case, coherence of the motion stimulus) when relating belief and performance (for example, their <xref ref-type="fig" rid="pone-0096511-g004">Fig. 4</xref>). However, as they compute the model's belief explicitly under the assumption of an unknown task difficulty, their performance predictions for different difficulties and its relation to the observed performance for these difficulties are in fact correct. The work in <xref ref-type="bibr" rid="pone.0096511-Kepecs1">[8]</xref>, in contrast, attempts to establish a direct relationship between the psychometric curve, conditional on task difficulty (the odor mixture ratio in their <xref ref-type="fig" rid="pone-0096511-g001">Fig 1c</xref>/d), and the decision maker's belief, as encoded by neurons in the orbifrontal cortex (their <xref ref-type="fig" rid="pone-0096511-g002">Fig. 2</xref>). As we have seen previously, this is the kind of situation in which belief and performance are not equal because performance is conditioned on task difficulty while task difficulty is unknown to the subject. This mismatch necessarily leads to miscalibration as illustrated in <xref ref-type="fig" rid="pone-0096511-g005">Fig 5b</xref>. Fortunately, the qualitative results of this particular study did not rely on a perfect match between belief and performance, but merely on a significant correlation between these two measures, which is likely to be true in their task, even if this correlation might be weak. A similar problem occurs in <xref ref-type="bibr" rid="pone.0096511-Grinband1">[11]</xref>, where the decision maker's confidence is directly derived from the psychometric curve (their <xref ref-type="fig" rid="pone-0096511-g002">Fig. 2a</xref>), again conditional on task difficulty (the width of the line that needs to be compared to a memorized reference), and is subsequently used as a parametric regressor in the analysis of functional magnet resonance imaging data. As we have demonstrated, there is no guarantee of a strong correlation between the psychometric curve and the decision maker's confidence, as for example demonstrated by a correlation coefficient below 0.35 between trial-by-trial belief and performance in <xref ref-type="fig" rid="pone-0096511-g005">Fig. 5b</xref>. Therefore, with this type of experiments. regressing performance against voxel activation only provides a weak test of whether an area is involved in encoding confidence. It is preferable to use instead a task in which the correlation between belief and performance is stronger, such as 2-AFC task in which subject knows the difficulty of the trial. Overall, these three examples demonstrate that the problems we have identified when relating belief and performance are not just obscure theoretical constructs, but occur in recent work in the neuroscience literature and have consequences for experimental design.</p>
<p>From the point-of-view of designing decision making models, our findings about the relation between belief and performance illustrate that models that aim to explain how humans and animals perform perceptual decision making should mostly focus on the encoded belief rather than on their performance. As long as they implement the correct generative model for the task, this belief will lead to the correct assessment of the model's task performance. For example, in both diffusion and race models, significant emphasis is put on expressions that describe the choice probability given some value of the hidden state, that is, the predicted performance <xref ref-type="bibr" rid="pone.0096511-Smith2">[47]</xref>. Instead, one should focus on the belief, which is the relevant quantity for the decision maker. A further advantage of this change of focus is that belief can be expressed analytically even for complex time-changing boundaries and arbitrary priors (see Methods: 2-AFC decision making with diffusion models), where no expressions for performance are known <xref ref-type="bibr" rid="pone.0096511-MorenoBote1">[27]</xref>. This simplifies the experimental validation of such models, as has been previously demonstrated in <xref ref-type="bibr" rid="pone.0096511-Drugowitsch1">[26]</xref>.</p>
<p>A further contribution of our work is to show that the decision maker's belief can in most cases only be measured on average, across many trials in which the decision maker's trial-by-trial belief might differ. The form of the average depends on one hand on the decision strategy of the decision maker (for example, diffusion model vs. race model) and on the other hand on the task setup. Being able to only control the latter, experimenters should thus attempt to avoid tasks in which measuring the decision maker's belief is important and trial-by-trial fluctuations around the measurable average can cause this measure to be only very weakly correlated to the belief in individual trials. This is, as we have established, to be expected in tasks with high-dimensional sufficient statistics of the decision maker's belief. Alternatively, the experimenter needs to commit to collecting data for a large number of trials to achieve a robust estimate of the decision maker's average belief despite strong trial-by-trial fluctuations around this average. A promising venue of research that would alleviate the problems of estimating belief from behavioral measurements is gathering more specific information about the decision maker's state by multiunit electrophysiological recordings of neural population activity.</p>
</sec><sec id="s4" sec-type="materials|methods">
<title>Materials and Methods</title>
<sec id="s4a">
<title>Decision-making framework</title>
<p>Here, we provide a brief description of the decision-making framework. For a more comprehensive discussion of its components, see Results. We assume that on each trial, the experimenter chooses a hidden state <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e320" xlink:type="simple"/></inline-formula> (e.g. the global direction of motion of a set of dots) according to the prior <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e321" xlink:type="simple"/></inline-formula>. The aim of the decision maker is to identify this hidden state by means of an observation <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e322" xlink:type="simple"/></inline-formula> (e.g. the motion energy in the display over a short time bin, or the neural activity in area MT) that relates to <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e323" xlink:type="simple"/></inline-formula> by the generative model <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e324" xlink:type="simple"/></inline-formula>, which is assumed to be known by the decision maker. In the following we show how both diffusion and race models can be described in this framework. Specifically, we derive the observation <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e325" xlink:type="simple"/></inline-formula> as the sufficient statistics of the posterior <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e326" xlink:type="simple"/></inline-formula>, and show that the decision time allows the experimenter to gain some limited information about <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e327" xlink:type="simple"/></inline-formula> without knowing its exact value.</p>
</sec><sec id="s4b">
<title>2-AFC decision making with diffusion models</title>
<p>In a diffusion model (DM), evidence about the hidden state <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e328" xlink:type="simple"/></inline-formula> is provided in each of a sequence of small time steps of size <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e329" xlink:type="simple"/></inline-formula> independently by the Gaussian momentary evidence <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e330" xlink:type="simple"/></inline-formula> with mean <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e331" xlink:type="simple"/></inline-formula> and variance <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e332" xlink:type="simple"/></inline-formula>. The mean rate <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e333" xlink:type="simple"/></inline-formula> is non-negative, <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e334" xlink:type="simple"/></inline-formula>, for <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e335" xlink:type="simple"/></inline-formula> and negative, <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e336" xlink:type="simple"/></inline-formula>, for <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e337" xlink:type="simple"/></inline-formula>. Its magnitude <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e338" xlink:type="simple"/></inline-formula> is a nuisance parameter that is uninformative about the hidden state, but determines the difficulty of the task.</p>
<p>We define the observation space <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e339" xlink:type="simple"/></inline-formula> by the sufficient statistics of the posterior belief given some sequence <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e340" xlink:type="simple"/></inline-formula> of momentary evidences from time <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e341" xlink:type="simple"/></inline-formula> to <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e342" xlink:type="simple"/></inline-formula> (such that <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e343" xlink:type="simple"/></inline-formula>) as follows. By Bayes' rule, and by the independence of the momentary evidences across time, the posterior <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e344" xlink:type="simple"/></inline-formula> is given by <disp-formula id="pone.0096511.e345"><graphic position="anchor" xlink:href="info:doi/10.1371/journal.pone.0096511.e345" xlink:type="simple"/><label>(14)</label></disp-formula>where all proportionalities are with respect to <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e346" xlink:type="simple"/></inline-formula>, and where we have used <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e347" xlink:type="simple"/></inline-formula> and <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e348" xlink:type="simple"/></inline-formula>. In the second-to-last line, the only dependency on the full trajectory not expressible through <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e349" xlink:type="simple"/></inline-formula> appears in the term in brackets, which is dropped in the last line, as it does not contain any <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e350" xlink:type="simple"/></inline-formula>-related terms. Thus, with <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e351" xlink:type="simple"/></inline-formula>, <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e352" xlink:type="simple"/></inline-formula> describes the location of a drifting and diffusing particle, <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e353" xlink:type="simple"/></inline-formula>. Here, <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e354" xlink:type="simple"/></inline-formula> is zero-mean Gaussian white noise with <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e355" xlink:type="simple"/></inline-formula> and <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e356" xlink:type="simple"/></inline-formula>, where <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e357" xlink:type="simple"/></inline-formula> is the Dirac delta function. This shows that, independently of the exact form of the prior <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e358" xlink:type="simple"/></inline-formula>, the posterior <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e359" xlink:type="simple"/></inline-formula> only depends on the current time <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e360" xlink:type="simple"/></inline-formula>, and the location <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e361" xlink:type="simple"/></inline-formula> of the drifting and diffusing particle at that time, rather than on the whole particle trajectory <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e362" xlink:type="simple"/></inline-formula>. Furthermore, by our definition, we have <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e363" xlink:type="simple"/></inline-formula> for all non-negative <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e364" xlink:type="simple"/></inline-formula>, such that <disp-formula id="pone.0096511.e365"><graphic position="anchor" xlink:href="info:doi/10.1371/journal.pone.0096511.e365" xlink:type="simple"/><label>(15)</label></disp-formula>which demonstrates that the decision maker's belief also depends only on <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e366" xlink:type="simple"/></inline-formula> and <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e367" xlink:type="simple"/></inline-formula>, for all possible priors <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e368" xlink:type="simple"/></inline-formula>. This holds even if the particle drifts in a bounded space with arbitrarily shaped boundaries <xref ref-type="bibr" rid="pone.0096511-Drugowitsch1">[26]</xref>–<xref ref-type="bibr" rid="pone.0096511-MorenoBote1">[27]</xref>. Thus, if we assume decisions to be triggered at the time-varying boundaries <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e369" xlink:type="simple"/></inline-formula> and <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e370" xlink:type="simple"/></inline-formula> with <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e371" xlink:type="simple"/></inline-formula> for all <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e372" xlink:type="simple"/></inline-formula>, and starting at <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e373" xlink:type="simple"/></inline-formula>, then we can define an observation by the belief's sufficient statistics at one of these boundaries. As a result, the observation is given by the pair <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e374" xlink:type="simple"/></inline-formula>, where <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e375" xlink:type="simple"/></inline-formula> is the decision time and <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e376" xlink:type="simple"/></inline-formula>. Furthermore, the set of possible observations is <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e377" xlink:type="simple"/></inline-formula>, where the last condition makes sure that the particle has not crossed either boundary before <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e378" xlink:type="simple"/></inline-formula>. Knowing the decision time <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e379" xlink:type="simple"/></inline-formula> thus restricts the set of possible observations to <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e380" xlink:type="simple"/></inline-formula>.</p>
</sec><sec id="s4c">
<title><inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e381" xlink:type="simple"/></inline-formula>-AFC decision making with race models</title>
<p>We assume a model with <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e382" xlink:type="simple"/></inline-formula> races, with race <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e383" xlink:type="simple"/></inline-formula> providing independent information by its associated drifting and diffusing particle <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e384" xlink:type="simple"/></inline-formula> with non-negative drift rate <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e385" xlink:type="simple"/></inline-formula>, and starting at <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e386" xlink:type="simple"/></inline-formula>. Here, the <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e387" xlink:type="simple"/></inline-formula>'s are uncorrelated unit-variance Gaussian white noises, such that <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e388" xlink:type="simple"/></inline-formula> and <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e389" xlink:type="simple"/></inline-formula>, where <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e390" xlink:type="simple"/></inline-formula> is the Dirac delta function, and <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e391" xlink:type="simple"/></inline-formula> is the Kronecker delta. The hidden state is associated with the fastest race, such that <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e392" xlink:type="simple"/></inline-formula> iff <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e393" xlink:type="simple"/></inline-formula>. The decision maker estimates this hidden state by forming a posterior over the drift rates given the full particle trajectory of all particles. As for the DM, we find this posterior by discretizing these particle trajectories into small time steps of size <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e394" xlink:type="simple"/></inline-formula>, such that in the <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e395" xlink:type="simple"/></inline-formula> th step, particle <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e396" xlink:type="simple"/></inline-formula> provides momentary evidence <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e397" xlink:type="simple"/></inline-formula>. If we assume to observe these trajectories from time <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e398" xlink:type="simple"/></inline-formula> to <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e399" xlink:type="simple"/></inline-formula>, the posterior over the drift rates becomes <disp-formula id="pone.0096511.e400"><graphic position="anchor" xlink:href="info:doi/10.1371/journal.pone.0096511.e400" xlink:type="simple"/><label>(16)</label></disp-formula>with all proportionalities with respect to the drift rates, where <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e401" xlink:type="simple"/></inline-formula>, and where we have used <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e402" xlink:type="simple"/></inline-formula> and <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e403" xlink:type="simple"/></inline-formula>. This shows that, as for the DM, this posterior depends only on time <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e404" xlink:type="simple"/></inline-formula> and the particle locations <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e405" xlink:type="simple"/></inline-formula> at this time, rather than the whole particle trajectory. From this posterior we find the hidden state posterior by <disp-formula id="pone.0096511.e406"><graphic position="anchor" xlink:href="info:doi/10.1371/journal.pone.0096511.e406" xlink:type="simple"/><label>(17)</label></disp-formula>which is again a function of only time and the current particle locations, thus forming the sufficient statistics of this belief. As before, the same sufficient statistics apply if the particle space is arbitrarily bounded.</p>
<p>A decision is made as soon as the first particle reaches a bound. If we assume that each race <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e407" xlink:type="simple"/></inline-formula> is upper-bounded independently by a time-varying boundary <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e408" xlink:type="simple"/></inline-formula> with <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e409" xlink:type="simple"/></inline-formula>, then the set of observations that describes the belief's sufficient statistics and that correspond to particle <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e410" xlink:type="simple"/></inline-formula> having reached the boundary first at time <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e411" xlink:type="simple"/></inline-formula> is <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e412" xlink:type="simple"/></inline-formula>, where the last condition again makes sure that no race has reached the boundary before <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e413" xlink:type="simple"/></inline-formula>. Thus, the set of observations that describe that a decision has been made at time <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e414" xlink:type="simple"/></inline-formula> is <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e415" xlink:type="simple"/></inline-formula>, that is, the set in which exactly one of the particles has reached the boundary at time <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e416" xlink:type="simple"/></inline-formula>. The set of all possible observations is thus given by <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e417" xlink:type="simple"/></inline-formula>. Importantly, an observation in either <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e418" xlink:type="simple"/></inline-formula>, <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e419" xlink:type="simple"/></inline-formula>, or <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e420" xlink:type="simple"/></inline-formula> does not only describe the state of the winning race, but also those of the losing races, as the belief depends on the state of all races. In Results we assume the same boundaries <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e421" xlink:type="simple"/></inline-formula> for all <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e422" xlink:type="simple"/></inline-formula> for convenience, but our formalism is also valid for boundary shapes that differ between races.</p>
</sec><sec id="s4d">
<title>An optimal decision strategy for the race model</title>
<p>Here we show that for a permutation-invariant prior <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e423" xlink:type="simple"/></inline-formula> on the drift rates, and the same bound, <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e424" xlink:type="simple"/></inline-formula>, on all races, a race model that chooses the option corresponding to the winning race corresponds to choosing the option that maximizes the posterior belief. The prior needs to be permutation-invariant in the sense that it needs to be invariant to swapping the values of any two drift rates. That is <disp-formula id="pone.0096511.e425"><graphic position="anchor" xlink:href="info:doi/10.1371/journal.pone.0096511.e425" xlink:type="simple"/><label>(18)</label></disp-formula>needs to hold for any two <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e426" xlink:type="simple"/></inline-formula>, where <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e427" xlink:type="simple"/></inline-formula> denote the random variable corresponding to the drift rate of race <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e428" xlink:type="simple"/></inline-formula>. In general, this can be achieved by defining the prior as a mixture of <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e429" xlink:type="simple"/></inline-formula> components (that is, the number possible swaps), each swapping two elements of a base distribution over <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e430" xlink:type="simple"/></inline-formula> random variables. A simpler, special case of this condition is a prior with <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e431" xlink:type="simple"/></inline-formula> mixture components that, for each component, assumes drift <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e432" xlink:type="simple"/></inline-formula> for all races except one, which instead features a drift of <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e433" xlink:type="simple"/></inline-formula>. The latter prior would correspond to the case where only a single race is informative about the correct option, while all the others are equally distractive.</p>
<p>To show optimality of choosing the option associated with the winning race, assume that race <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e434" xlink:type="simple"/></inline-formula> was the first to have reached the boundary <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e435" xlink:type="simple"/></inline-formula> at time <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e436" xlink:type="simple"/></inline-formula>. We demonstrate that, under these circumstances, the posterior belief of <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e437" xlink:type="simple"/></inline-formula> according to Eq. (17) is at least as large as for any other <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e438" xlink:type="simple"/></inline-formula> where <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e439" xlink:type="simple"/></inline-formula>. Choosing some arbitrary <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e440" xlink:type="simple"/></inline-formula>, we define for the observed <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e441" xlink:type="simple"/></inline-formula>, <disp-formula id="pone.0096511.e442"><graphic position="anchor" xlink:href="info:doi/10.1371/journal.pone.0096511.e442" xlink:type="simple"/><label>(19)</label></disp-formula>which, due to the permutation-invariant prior, is a non-negative symmetric function, that is <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e443" xlink:type="simple"/></inline-formula> and <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e444" xlink:type="simple"/></inline-formula>. This allows us to write the beliefs of <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e445" xlink:type="simple"/></inline-formula> and <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e446" xlink:type="simple"/></inline-formula> by Eqs. (16) and (17) as <disp-formula id="pone.0096511.e447"><graphic position="anchor" xlink:href="info:doi/10.1371/journal.pone.0096511.e447" xlink:type="simple"/><label>(20)</label></disp-formula>Thus, in order to satisfy <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e448" xlink:type="simple"/></inline-formula>, we need to have <disp-formula id="pone.0096511.e449"><graphic position="anchor" xlink:href="info:doi/10.1371/journal.pone.0096511.e449" xlink:type="simple"/><label>(21)</label></disp-formula>where we have substituted <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e450" xlink:type="simple"/></inline-formula> and <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e451" xlink:type="simple"/></inline-formula> on the left-hand side, and <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e452" xlink:type="simple"/></inline-formula> and <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e453" xlink:type="simple"/></inline-formula> on the right-hand side. Due to the non-negativity of <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e454" xlink:type="simple"/></inline-formula> and the strictly increasing and non-negative exponential, Eq. (21) is satisfied if <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e455" xlink:type="simple"/></inline-formula> for <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e456" xlink:type="simple"/></inline-formula> (due to the upper limit of the inner integral) and <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e457" xlink:type="simple"/></inline-formula> (race <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e458" xlink:type="simple"/></inline-formula> is winner, such that <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e459" xlink:type="simple"/></inline-formula> and <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e460" xlink:type="simple"/></inline-formula>). This is easily shown by using <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e461" xlink:type="simple"/></inline-formula>, such that this inequality can be written as <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e462" xlink:type="simple"/></inline-formula>, which, due to <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e463" xlink:type="simple"/></inline-formula> and <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e464" xlink:type="simple"/></inline-formula>, is always satisfied. As <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e465" xlink:type="simple"/></inline-formula> was arbitrarily chosen, it holds for all <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e466" xlink:type="simple"/></inline-formula>, such that choosing the option corresponding to the winning race guarantees that no other choice would have led to a higher belief of being correct.</p>
</sec><sec id="s4e">
<title>Equivalence of average and trial-by-trial belief</title>
<p>After observing <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e467" xlink:type="simple"/></inline-formula> in a given trial, the decision maker commits to decision <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e468" xlink:type="simple"/></inline-formula>, where <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e469" xlink:type="simple"/></inline-formula>, and holds belief <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e470" xlink:type="simple"/></inline-formula>. Knowing only that <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e471" xlink:type="simple"/></inline-formula>, the experimenter can measure the average belief <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e472" xlink:type="simple"/></inline-formula> across multiple trials. As we discuss in more detail in Results, the relation between the decision maker's trial-by-trial belief and the average belief measured by the experimenter is <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e473" xlink:type="simple"/></inline-formula>, as given by Eq. (8). This relation states that the average belief is the trial-by-trial belief averaged over all trials in which option <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e474" xlink:type="simple"/></inline-formula> is chosen, and in which the observation <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e475" xlink:type="simple"/></inline-formula> conforms to <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e476" xlink:type="simple"/></inline-formula>. Clearly, if the set <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e477" xlink:type="simple"/></inline-formula> only holds a single <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e478" xlink:type="simple"/></inline-formula>, such that <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e479" xlink:type="simple"/></inline-formula> is only non-zero for this one <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e480" xlink:type="simple"/></inline-formula>, then average belief and trial-by-trial belief are equivalent. Here we consider a slightly more general condition based on the minimal sufficient statistics of the trial-by-trial belief.</p>
<p>Intuitively, a necessary and sufficient condition for the equivalence of average and trial-by-trial belief is that, for all trials that we average over, the observation <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e481" xlink:type="simple"/></inline-formula> needs to lead to the same trial-by-trial belief. Thus, while it is permissible for <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e482" xlink:type="simple"/></inline-formula> to vary across trials, its contribution to the trial-by-trial belief needs to remain constant. This contribution is formalized as the minimal sufficient statistic <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e483" xlink:type="simple"/></inline-formula> of <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e484" xlink:type="simple"/></inline-formula> with respect to <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e485" xlink:type="simple"/></inline-formula>, such that <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e486" xlink:type="simple"/></inline-formula>. If we have two observations, <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e487" xlink:type="simple"/></inline-formula> and <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e488" xlink:type="simple"/></inline-formula>, for which <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e489" xlink:type="simple"/></inline-formula>, it is by the definition of minimal sufficient statistics guaranteed that <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e490" xlink:type="simple"/></inline-formula>. In contrast, for two observations for which <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e491" xlink:type="simple"/></inline-formula> we have <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e492" xlink:type="simple"/></inline-formula> even if <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e493" xlink:type="simple"/></inline-formula>. Thus, a necessary and sufficient condition for the equivalence of average and trial-by-trial belief is that for all <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e494" xlink:type="simple"/></inline-formula> such that <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e495" xlink:type="simple"/></inline-formula> we need to guarantee that <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e496" xlink:type="simple"/></inline-formula> is the same constant. This condition is sufficient, because if it holds, then <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e497" xlink:type="simple"/></inline-formula> will be the same for all trials that we average over, resulting in an equivalent average belief. It is necessary, because if <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e498" xlink:type="simple"/></inline-formula> differs for at least one <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e499" xlink:type="simple"/></inline-formula>, then we will average over different trial-by-trial beliefs.</p>
<p>To relate this condition to DMs and race models, let us consider their minimal sufficient statistics. In the case of diffusion models, these statistics are particle location and time, <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e500" xlink:type="simple"/></inline-formula>. Thus, as <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e501" xlink:type="simple"/></inline-formula> and <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e502" xlink:type="simple"/></inline-formula> imply that a particular boundary has been reached at a known time, both <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e503" xlink:type="simple"/></inline-formula> and <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e504" xlink:type="simple"/></inline-formula> are guaranteed to be uniquely determined, such that <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e505" xlink:type="simple"/></inline-formula> is constant under these conditions. As a result, average belief and trial-to-trial belief are equivalent. In <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e506" xlink:type="simple"/></inline-formula>-race models, on the other hand, the sufficient statistics are the state of all races and time, <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e507" xlink:type="simple"/></inline-formula>. In this case, <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e508" xlink:type="simple"/></inline-formula> and <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e509" xlink:type="simple"/></inline-formula> restrict time and the state of the winning race, but not that of the other races (other than them being below the boundary), such that <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e510" xlink:type="simple"/></inline-formula> can change between trials. As a consequence, the trial-by-trial belief will fluctuate around the average belief.</p>
</sec><sec id="s4f">
<title>Minimal sufficient statistics in an orientation categorization task</title>
<p>To show that the dimensionality of the minimal sufficient statistics does not necessarily grow with the number of options available to the decision maker, consider the following task. Assume a set of <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e511" xlink:type="simple"/></inline-formula> orientations, <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e512" xlink:type="simple"/></inline-formula>, on a half-circle, <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e513" xlink:type="simple"/></inline-formula>, with the <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e514" xlink:type="simple"/></inline-formula>th orientation corresponding to hidden state <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e515" xlink:type="simple"/></inline-formula>. In each trial, the experimenter picks a hidden state <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e516" xlink:type="simple"/></inline-formula> which is used to generate an oriented stimulus with orientation <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e517" xlink:type="simple"/></inline-formula> by drawing <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e518" xlink:type="simple"/></inline-formula> from a von Mises distribution with mean <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e519" xlink:type="simple"/></inline-formula> and concentration <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e520" xlink:type="simple"/></inline-formula>. The decision maker perceives this orientation with some additional sensory noise, such that the likelihood of the decision maker's observation <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e521" xlink:type="simple"/></inline-formula> is <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e522" xlink:type="simple"/></inline-formula> with reduced concentration <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e523" xlink:type="simple"/></inline-formula>, and thus lower precision. Assuming a uniform prior, <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e524" xlink:type="simple"/></inline-formula>, and that the decision maker has learned concentration <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e525" xlink:type="simple"/></inline-formula> over past trials, her belief over the hidden states follows from Bayes' rule and the definition of the von Mises distribution, and is given by <disp-formula id="pone.0096511.e526"><graphic position="anchor" xlink:href="info:doi/10.1371/journal.pone.0096511.e526" xlink:type="simple"/><label>(22)</label></disp-formula>This shows that, independent of the decision maker's decision function <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e527" xlink:type="simple"/></inline-formula>, a minimal sufficient statistic of her trial-by-trial belief is the observation, <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e528" xlink:type="simple"/></inline-formula>, whose dimensionality is always one, independent of the number <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e529" xlink:type="simple"/></inline-formula> of possible options to choose from.</p>
</sec><sec id="s4g">
<title>Generating <xref ref-type="fig" rid="pone-0096511-g003">Figures 3</xref> and <xref ref-type="fig" rid="pone-0096511-g004">4</xref></title>
<p>Here we explain how we simulated the diffusion model (DM) and 2-race model to generate <xref ref-type="fig" rid="pone-0096511-g003">Figs. 3</xref> and <xref ref-type="fig" rid="pone-0096511-g004">4</xref>. For both model types, we determined choices and decision times by the decision models, and computed the reaction times by adding a fixed non-decision time of 250 ms to each decision time. All simulations were performed in 1 ms time steps up to a maximum of 2 s, after which the simulation was aborted.</p>
<p>For the DM, we assumed <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e530" xlink:type="simple"/></inline-formula> for <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e531" xlink:type="simple"/></inline-formula> and <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e532" xlink:type="simple"/></inline-formula> for <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e533" xlink:type="simple"/></inline-formula>, with <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e534" xlink:type="simple"/></inline-formula>. The upper and lower boundaries were time-varying and symmetric around zero, defined by <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e535" xlink:type="simple"/></inline-formula> and <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e536" xlink:type="simple"/></inline-formula> respectively. We have chosen a time-varying boundary to have the belief at the boundary to depend on time. If we had been using a time-invariant boundary instead, this time-dependency of the belief would vanish. Given this setup, we found the decision maker's belief when reaching the upper boundary and thus choosing <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e537" xlink:type="simple"/></inline-formula> by Eq. (14), resulting in <disp-formula id="pone.0096511.e538"><graphic position="anchor" xlink:href="info:doi/10.1371/journal.pone.0096511.e538" xlink:type="simple"/><label>(23)</label></disp-formula>where <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e539" xlink:type="simple"/></inline-formula>, and <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e540" xlink:type="simple"/></inline-formula> is the normalization constant. We find <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e541" xlink:type="simple"/></inline-formula> by solving <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e542" xlink:type="simple"/></inline-formula>, which, when substituted into Eq. (23) a re-arranging the terms, results in the final belief <disp-formula id="pone.0096511.e543"><graphic position="anchor" xlink:href="info:doi/10.1371/journal.pone.0096511.e543" xlink:type="simple"/><label>(24)</label></disp-formula>For the uniform prior case in <xref ref-type="fig" rid="pone-0096511-g003">Fig. 3a</xref> we generated 10000 trials for each <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e544" xlink:type="simple"/></inline-formula> and <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e545" xlink:type="simple"/></inline-formula>, simulating <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e546" xlink:type="simple"/></inline-formula> in small time steps until either boundary was reached. We then binned trials by decision time in bins of 250 ms from 250 ms to 1500 ms. To compute performance for each bin we randomly picked 500 trials from this bin in which <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e547" xlink:type="simple"/></inline-formula>, and computed the fraction of times that the upper boundary was reached. Additionally, we plotted the belief for 10 randomly chosen trials from this bin in which this upper boundary was reached.</p>
<p>For the non-uniform prior case in <xref ref-type="fig" rid="pone-0096511-g003">Figs. 3b</xref> and <xref ref-type="fig" rid="pone-0096511-g004">4a</xref> we chose <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e548" xlink:type="simple"/></inline-formula>. We then generated 10000 trials with <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e549" xlink:type="simple"/></inline-formula> and (to conform to the prior) 5384 trials with <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e550" xlink:type="simple"/></inline-formula> by again simulating <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e551" xlink:type="simple"/></inline-formula> in small time steps. Due to the non-uniform prior, all trajectories reaching the upper boundary caused choice 1, but only trajectories that reached the lower boundary below <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e552" xlink:type="simple"/></inline-formula> caused decision 2, and decision 1 otherwise. This strategy arises because the belief at low boundaries is close to <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e553" xlink:type="simple"/></inline-formula>. In these cases, the prior might provide more evidence than the likelihood, which might cause a reversal of the decision if prior and likelihood provide evidence for opposing options. Performance and belief were again computed/selected as for the uniform prior case. We computed the estimated belief in <xref ref-type="fig" rid="pone-0096511-g003">Fig. 3b</xref> by the fraction of correct choices among 500 trials per bin in which option 1 was chosen.</p>
<p>For the 2-race model in <xref ref-type="fig" rid="pone-0096511-g003">Figs. 3c</xref> and <xref ref-type="fig" rid="pone-0096511-g004">4b</xref> we chose <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e554" xlink:type="simple"/></inline-formula> for <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e555" xlink:type="simple"/></inline-formula> and <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e556" xlink:type="simple"/></inline-formula> for <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e557" xlink:type="simple"/></inline-formula>, with <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e558" xlink:type="simple"/></inline-formula>. We used the boundary <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e559" xlink:type="simple"/></inline-formula>, that varied over time but not between races. Given this setup, the decision maker's belief when race 1 is the winning race follows from Eq. (16) and is given by <disp-formula id="pone.0096511.e560"><graphic position="anchor" xlink:href="info:doi/10.1371/journal.pone.0096511.e560" xlink:type="simple"/><label>(25)</label></disp-formula>which is a function of both the bound height and the state of the second race. We simulated 10000 trials for each <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e561" xlink:type="simple"/></inline-formula> in small time steps, and binned trials by decision time into 250 ms bins from 250 ms to 1750 ms. Performance, trial-by-trial belief, and estimated belief were computed and plotted as for the DM.</p>
</sec><sec id="s4h">
<title>Computing belief in a drift diffusion model with varying difficulty (<xref ref-type="fig" rid="pone-0096511-g005">Figure 5</xref>)</title>
<p>We generated <xref ref-type="fig" rid="pone-0096511-g005">Fig. 5</xref> by assuming a decision making diffusion model with diffusion variance <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e562" xlink:type="simple"/></inline-formula>, and time-invariant bounds at <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e563" xlink:type="simple"/></inline-formula>. Note that with this choice of diffusion variance, the drift and <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e564" xlink:type="simple"/></inline-formula> are measured in units of <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e565" xlink:type="simple"/></inline-formula>. The drift rate was constant within a trial and was chosen across trials to roughly follow <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e566" xlink:type="simple"/></inline-formula> with <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e567" xlink:type="simple"/></inline-formula> (in units of <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e568" xlink:type="simple"/></inline-formula>). Specifically, we used a point-wise drift rate prior corresponding to a uniform distribution over nine different drift rates, where <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e569" xlink:type="simple"/></inline-formula> corresponds to the 10<sup>th</sup> percentile of <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e570" xlink:type="simple"/></inline-formula>, <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e571" xlink:type="simple"/></inline-formula> corresponding to the 20<sup>th</sup> percentile, and so on, up to <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e572" xlink:type="simple"/></inline-formula> for the 90<sup>th</sup> percentile (<xref ref-type="fig" rid="pone-0096511-g005">Fig 5a</xref>). Reaction times and choice probability were computed analytically using standard results for bounded diffusion models <xref ref-type="bibr" rid="pone.0096511-Cox1">[48]</xref>–<xref ref-type="bibr" rid="pone.0096511-Palmer1">[49]</xref>.</p>
<p>For <xref ref-type="fig" rid="pone-0096511-g005">Fig 5b</xref> we computed the decision maker's belief under two different assumptions. First, we assumed exact knowledge of the correct point-wise prior, <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e573" xlink:type="simple"/></inline-formula> for <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e574" xlink:type="simple"/></inline-formula>, for which the posterior drift rate given that the particle reached the upper or lower bound <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e575" xlink:type="simple"/></inline-formula> at time <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e576" xlink:type="simple"/></inline-formula> follows Eq. (14), and results in <xref ref-type="bibr" rid="pone.0096511-Drugowitsch1">[26]</xref> <disp-formula id="pone.0096511.e577"><graphic position="anchor" xlink:href="info:doi/10.1371/journal.pone.0096511.e577" xlink:type="simple"/><label>(26)</label></disp-formula>To find the belief we split the prior mass of <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e578" xlink:type="simple"/></inline-formula> uniformly between <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e579" xlink:type="simple"/></inline-formula> and <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e580" xlink:type="simple"/></inline-formula> by replacing it by <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e581" xlink:type="simple"/></inline-formula> and <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e582" xlink:type="simple"/></inline-formula> with prior masses <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e583" xlink:type="simple"/></inline-formula>, and assign all positive drift rates <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e584" xlink:type="simple"/></inline-formula> and <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e585" xlink:type="simple"/></inline-formula> to hidden state <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e586" xlink:type="simple"/></inline-formula>, while the remaining drift rates correspond to hidden state <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e587" xlink:type="simple"/></inline-formula>. When hitting the upper bound and choosing <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e588" xlink:type="simple"/></inline-formula>, this results in the belief <disp-formula id="pone.0096511.e589"><graphic position="anchor" xlink:href="info:doi/10.1371/journal.pone.0096511.e589" xlink:type="simple"/><label>(27)</label></disp-formula>Due to symmetry of prior and task, the same equation holds for the belief of <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e590" xlink:type="simple"/></inline-formula> when hitting the lower bound and choosing <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e591" xlink:type="simple"/></inline-formula>. This is the optimal belief the decision maker can hold in this task. Second, we computed the belief based on the assumption that, instead of the correct, point-wise prior, the decision maker assumes a Gaussian zero-mean prior, <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e592" xlink:type="simple"/></inline-formula> whose variance <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e593" xlink:type="simple"/></inline-formula> might differ from <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e594" xlink:type="simple"/></inline-formula>. This allowed us to simulate cases in which the decision maker uses an incorrect prior. With this prior, the belief follows again Eqs. (14) and (15), resulting in <xref ref-type="bibr" rid="pone.0096511-Drugowitsch1">[26]</xref> <disp-formula id="pone.0096511.e595"><graphic position="anchor" xlink:href="info:doi/10.1371/journal.pone.0096511.e595" xlink:type="simple"/><label>(28)</label></disp-formula>where <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e596" xlink:type="simple"/></inline-formula> is the standard cumulative Gaussian. To find the average belief per drift rate, as shown in <xref ref-type="fig" rid="pone-0096511-g005">Fig. 5b</xref>, we numerically computed the reaction time distribution <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e597" xlink:type="simple"/></inline-formula> for each <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e598" xlink:type="simple"/></inline-formula> in steps of 1 ms up to <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e599" xlink:type="simple"/></inline-formula> as the solution of a Volterra integral equation of the second kind <xref ref-type="bibr" rid="pone.0096511-Smith2">[47]</xref>. Based on this, we computed the average belief for both the point-wise and the Gaussian prior (with <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e600" xlink:type="simple"/></inline-formula>) by numerically evaluating the integral <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e601" xlink:type="simple"/></inline-formula>. The standard deviation of the belief for the point-wise prior was similarly evaluated by numerical integration based on these reaction-time distributions.</p>
<p>The calibration curves in <xref ref-type="fig" rid="pone-0096511-g005">Fig. 5c</xref> were found as follows. Conditional on the absolute drift rate, the probability of performing correct choices is given by <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e602" xlink:type="simple"/></inline-formula> and is thus independent of the reaction time <xref ref-type="bibr" rid="pone.0096511-Cox1">[48]</xref>. In contrast, the belief (Eq. (27)) depends on the reaction time, such that, for a fixed drift rate it will vary across trials even if the probability of choosing the correction option does not. As a result, the calibration curves conditioned on the drift rate, which are given by the function <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e603" xlink:type="simple"/></inline-formula> of belief <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e604" xlink:type="simple"/></inline-formula>, are independent of this belief and thus flat (<xref ref-type="fig" rid="pone-0096511-g005">Fig. 5c</xref>). This does not hold anymore as soon as we consider the average calibration curve, as given by <disp-formula id="pone.0096511.e605"><graphic position="anchor" xlink:href="info:doi/10.1371/journal.pone.0096511.e605" xlink:type="simple"/><label>(29)</label></disp-formula>In this case, the probability of making a correct choice depends on the reaction time, as the distribution of these reaction times differs for different drift rates. This becomes particularly clear when expanding the choice probability to give</p>
<p><disp-formula id="pone.0096511.e606"><graphic position="anchor" xlink:href="info:doi/10.1371/journal.pone.0096511.e606" xlink:type="simple"/><label>(30)</label></disp-formula>where the fraction inside the sum results from Bayes rule applied to <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e607" xlink:type="simple"/></inline-formula>. In the above sum, the first term is the known and time-invariant choice probability for a fixed drift rate. The time-dependence is introduced in the second term which is proportional to the probability of reaction time <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e608" xlink:type="simple"/></inline-formula> for drift rate <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e609" xlink:type="simple"/></inline-formula> and hence a function of both variables. To evaluate Eq. (29) for a given <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e610" xlink:type="simple"/></inline-formula> we can utilize the fact that the belief is monotonic in time, such that <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e611" xlink:type="simple"/></inline-formula> is a Dirac-delta function at the <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e612" xlink:type="simple"/></inline-formula> where <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e613" xlink:type="simple"/></inline-formula>. Thus, Eq. (29) results in Eq. (30) evaluated at this time <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e614" xlink:type="simple"/></inline-formula>, which we find by using the numerical reaction time distributions for fixed drift rates while being again careful about splitting the mass of <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e615" xlink:type="simple"/></inline-formula> in half. Evaluated for each valid <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e616" xlink:type="simple"/></inline-formula> results in the average calibration curve shown in <xref ref-type="fig" rid="pone-0096511-g005">Fig. 5c</xref>. In Results (see Eq. (11)) we explain why this average curve follows the identity line.</p>
<p>To simulate the calibration curves for subjects using an incorrect prior <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e617" xlink:type="simple"/></inline-formula>, as shown in <xref ref-type="fig" rid="pone-0096511-g005">Fig. 5d</xref>, we used the same procedure as to compute the average calibration curve in <xref ref-type="fig" rid="pone-0096511-g005">Fig. 5c</xref>. However, rather than using the correct point-wise prior to compute the belief, we assumed the decision maker to utilize a Gaussian prior with different standard deviations. Furthermore, we assumed the actual drift rates to follow a Gaussian prior such that in Eq. (30) the sum turns into an integral that we solved numerically. Specifically, to simulate a too-wide prior, we assumed the actual prior to be a zero-mean Gaussian with standard deviation <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e618" xlink:type="simple"/></inline-formula>, while the decision maker assumed <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e619" xlink:type="simple"/></inline-formula>. For the too narrow prior we used the actual prior width <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e620" xlink:type="simple"/></inline-formula> while setting the assumed prior width to <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e621" xlink:type="simple"/></inline-formula>.</p>
</sec><sec id="s4i">
<title>Modeling miscalibrations by the decision variable partition model</title>
<p>The decision variable partition model <xref ref-type="bibr" rid="pone.0096511-Ferrell1">[32]</xref>–<xref ref-type="bibr" rid="pone.0096511-Ferrell2">[33]</xref> is a popular model based on signal detection theory to explain various types of miscalibration. In this model, the decision maker observes two random variables, <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e622" xlink:type="simple"/></inline-formula> and <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e623" xlink:type="simple"/></inline-formula> (for example, two weights that need to be compared), where one is drawn from the “correct”, and the other from the “incorrect” distribution and the aim of the decision maker is to identify the one associated with the correct distribution. We formalize this by <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e624" xlink:type="simple"/></inline-formula> and <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e625" xlink:type="simple"/></inline-formula> for <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e626" xlink:type="simple"/></inline-formula>, and <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e627" xlink:type="simple"/></inline-formula> and <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e628" xlink:type="simple"/></inline-formula> with flipped means for <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e629" xlink:type="simple"/></inline-formula>. Then, it is easy to show that the hidden state posterior upon observing <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e630" xlink:type="simple"/></inline-formula> and <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e631" xlink:type="simple"/></inline-formula> is given by <disp-formula id="pone.0096511.e632"><graphic position="anchor" xlink:href="info:doi/10.1371/journal.pone.0096511.e632" xlink:type="simple"/><label>(31)</label></disp-formula>where <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e633" xlink:type="simple"/></inline-formula> determines the difficulty of the task. Thus, the decision rule that maximizes this posterior is to choose <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e634" xlink:type="simple"/></inline-formula> if <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e635" xlink:type="simple"/></inline-formula> and <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e636" xlink:type="simple"/></inline-formula> otherwise. This leads to the optimal belief <disp-formula id="pone.0096511.e637"><graphic position="anchor" xlink:href="info:doi/10.1371/journal.pone.0096511.e637" xlink:type="simple"/><label>(32)</label></disp-formula>which is a monotonically increasing function of <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e638" xlink:type="simple"/></inline-formula> (a larger difference in perceived weights increases the confidence). Instead of using the optimal belief according to Eq. (32), the decision variable partition model proposes to partitioning the space of <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e639" xlink:type="simple"/></inline-formula> into arbitrarily chosen bins and to assign each of these bins a different confidence rating. This way it is able to capture any deviation from perfect calibration, as long as the decision maker's performance grows monotonically with belief. Furthermore, it captures the hard-easy effect by leaving the partitioning unchanged for different task difficulties while the optimal belief would require adjusting <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pone.0096511.e640" xlink:type="simple"/></inline-formula> in Eq. (32) accordingly.</p>
</sec></sec></body>
<back>
<ack>
<p>We would like to thank Peter Latham for comments on early version of this manuscript and Simon Barthelme for advice on the human confidence calibration literature.</p>
</ack>
<ref-list>
<title>References</title>
<ref id="pone.0096511-Beck1"><label>1</label>
<mixed-citation publication-type="journal" xlink:type="simple"><name name-style="western"><surname>Beck</surname><given-names>JM</given-names></name>, <name name-style="western"><surname>Ma</surname><given-names>WJ</given-names></name>, <name name-style="western"><surname>Kiani</surname><given-names>R</given-names></name>, <name name-style="western"><surname>Hanks</surname><given-names>T</given-names></name>, <name name-style="western"><surname>Churchland</surname><given-names>AK</given-names></name>, <etal>et al</etal>. (<year>2008</year>) <article-title>Probabilistic population codes for Bayesian decision making</article-title>. <source>Neuron</source> <volume>60</volume>: <fpage>1142</fpage>–<lpage>1152</lpage>.</mixed-citation>
</ref>
<ref id="pone.0096511-Alais1"><label>2</label>
<mixed-citation publication-type="journal" xlink:type="simple"><name name-style="western"><surname>Alais</surname><given-names>D</given-names></name>, <name name-style="western"><surname>Burr</surname><given-names>D</given-names></name> (<year>2004</year>) <article-title>The ventriloquist effect results from near-optimal bimodal integration</article-title>. <source>Curr Biol</source> <volume>14</volume>: <fpage>257</fpage>–<lpage>262</lpage>.</mixed-citation>
</ref>
<ref id="pone.0096511-Ernst1"><label>3</label>
<mixed-citation publication-type="journal" xlink:type="simple"><name name-style="western"><surname>Ernst</surname><given-names>MO</given-names></name>, <name name-style="western"><surname>Banks</surname><given-names>MS</given-names></name> (<year>2002</year>) <article-title>Humans integrate visual and haptic information in a statistically optimal fashion</article-title>. <source>Nature</source> <volume>415</volume>: <fpage>429</fpage>–<lpage>433</lpage>.</mixed-citation>
</ref>
<ref id="pone.0096511-Fetsch1"><label>4</label>
<mixed-citation publication-type="journal" xlink:type="simple"><name name-style="western"><surname>Fetsch</surname><given-names>CR</given-names></name>, <name name-style="western"><surname>Pouget</surname><given-names>A</given-names></name>, <name name-style="western"><surname>Deangelis</surname><given-names>GC</given-names></name>, <name name-style="western"><surname>Angelaki</surname><given-names>DE</given-names></name> (<year>2011</year>) <article-title>Neural correlates of reliability-based cue weighting during multisensory integration</article-title>. <source>Nat Neurosci</source> <volume>15</volume>: <fpage>146</fpage>–<lpage>154</lpage>.</mixed-citation>
</ref>
<ref id="pone.0096511-Kiani1"><label>5</label>
<mixed-citation publication-type="journal" xlink:type="simple"><name name-style="western"><surname>Kiani</surname><given-names>R</given-names></name>, <name name-style="western"><surname>Shadlen</surname><given-names>MN</given-names></name> (<year>2009</year>) <article-title>Representation of confidence associated with a decision by neurons in the parietal cortex</article-title>. <source>Science</source> <volume>324</volume>: <fpage>759</fpage>–<lpage>764</lpage>.</mixed-citation>
</ref>
<ref id="pone.0096511-vanBeers1"><label>6</label>
<mixed-citation publication-type="journal" xlink:type="simple"><name name-style="western"><surname>van Beers</surname><given-names>RJ</given-names></name>, <name name-style="western"><surname>Sittig</surname><given-names>AC</given-names></name>, <name name-style="western"><surname>Denier van der Gon</surname><given-names>JJ</given-names></name> (<year>1996</year>) <article-title>How humans combine simultaneous proprioceptive and visual position information</article-title>. <source>Exp Brain Res</source> <volume>111</volume>: <fpage>253</fpage>–<lpage>261</lpage>.</mixed-citation>
</ref>
<ref id="pone.0096511-Vickers1"><label>7</label>
<mixed-citation publication-type="book" xlink:type="simple">Vickers D (1979) Decision processes in visual perception. New York; London: Academic Press. <volume>vi</volume>: , 406 p. p.</mixed-citation>
</ref>
<ref id="pone.0096511-Kepecs1"><label>8</label>
<mixed-citation publication-type="journal" xlink:type="simple"><name name-style="western"><surname>Kepecs</surname><given-names>A</given-names></name>, <name name-style="western"><surname>Uchida</surname><given-names>N</given-names></name>, <name name-style="western"><surname>Zariwala</surname><given-names>HA</given-names></name>, <name name-style="western"><surname>Mainen</surname><given-names>ZF</given-names></name> (<year>2008</year>) <article-title>Neural correlates, computation and behavioural impact of decision confidence</article-title>. <source>Nature</source> <volume>455</volume>: <fpage>227</fpage>–<lpage>231</lpage>.</mixed-citation>
</ref>
<ref id="pone.0096511-Metcalfe1"><label>9</label>
<mixed-citation publication-type="book" xlink:type="simple">Metcalfe J, Shimamura AP (1994) Metacognition: knowing about knowing. Cambridge, Mass; London: MIT Press. <volume>xiii</volume>: , 334 p. p.</mixed-citation>
</ref>
<ref id="pone.0096511-Smith1"><label>10</label>
<mixed-citation publication-type="journal" xlink:type="simple"><name name-style="western"><surname>Smith</surname><given-names>JD</given-names></name>, <name name-style="western"><surname>Shields</surname><given-names>WE</given-names></name>, <name name-style="western"><surname>Washburn</surname><given-names>DA</given-names></name> (<year>2003</year>) <article-title>The comparative psychology of uncertainty monitoring and metacognition</article-title>. <source>Behav Brain Sci</source> <volume>26</volume>: <fpage>317</fpage>–<lpage>339</lpage> discussion 340–373.</mixed-citation>
</ref>
<ref id="pone.0096511-Grinband1"><label>11</label>
<mixed-citation publication-type="journal" xlink:type="simple"><name name-style="western"><surname>Grinband</surname><given-names>J</given-names></name>, <name name-style="western"><surname>Hirsch</surname><given-names>J</given-names></name>, <name name-style="western"><surname>Ferrera</surname><given-names>VP</given-names></name> (<year>2006</year>) <article-title>A neural representation of categorization uncertainty in the human brain</article-title>. <source>Neuron</source> <volume>49</volume>: <fpage>757</fpage>–<lpage>763</lpage>.</mixed-citation>
</ref>
<ref id="pone.0096511-Gigerenzer1"><label>12</label>
<mixed-citation publication-type="journal" xlink:type="simple"><name name-style="western"><surname>Gigerenzer</surname><given-names>G</given-names></name>, <name name-style="western"><surname>Hoffrage</surname><given-names>U</given-names></name>, <name name-style="western"><surname>Kleinbolting</surname><given-names>H</given-names></name> (<year>1991</year>) <article-title>Probabilistic mental models: a Brunswikian theory of confidence</article-title>. <source>Psychol Rev</source> <volume>98</volume>: <fpage>506</fpage>–<lpage>528</lpage>.</mixed-citation>
</ref>
<ref id="pone.0096511-Ratcliff1"><label>13</label>
<mixed-citation publication-type="journal" xlink:type="simple"><name name-style="western"><surname>Ratcliff</surname><given-names>R</given-names></name>, <name name-style="western"><surname>Starns</surname><given-names>JJ</given-names></name> (<year>2009</year>) <article-title>Modeling confidence and response time in recognition memory</article-title>. <source>Psychol Rev</source> <volume>116</volume>: <fpage>59</fpage>–<lpage>83</lpage>.</mixed-citation>
</ref>
<ref id="pone.0096511-DeMartino1"><label>14</label>
<mixed-citation publication-type="journal" xlink:type="simple"><name name-style="western"><surname>De Martino</surname><given-names>B</given-names></name>, <name name-style="western"><surname>Fleming</surname><given-names>SM</given-names></name>, <name name-style="western"><surname>Garrett</surname><given-names>N</given-names></name>, <name name-style="western"><surname>Dolan</surname><given-names>RJ</given-names></name> (<year>2013</year>) <article-title>Confidence in value-based choice</article-title>. <source>Nat Neurosci</source> <volume>16</volume>: <fpage>105</fpage>–<lpage>110</lpage>.</mixed-citation>
</ref>
<ref id="pone.0096511-Pleskac1"><label>15</label>
<mixed-citation publication-type="journal" xlink:type="simple"><name name-style="western"><surname>Pleskac</surname><given-names>TJ</given-names></name>, <name name-style="western"><surname>Busemeyer</surname><given-names>JR</given-names></name> (<year>2010</year>) <article-title>Two-stage dynamic signal detection: a theory of choice, decision time, and confidence</article-title>. <source>Psychol Rev</source> <volume>117</volume>: <fpage>864</fpage>–<lpage>901</lpage>.</mixed-citation>
</ref>
<ref id="pone.0096511-Juslin1"><label>16</label>
<mixed-citation publication-type="journal" xlink:type="simple"><name name-style="western"><surname>Juslin</surname><given-names>P</given-names></name>, <name name-style="western"><surname>Olsson</surname><given-names>H</given-names></name>, <name name-style="western"><surname>Bjorkman</surname><given-names>M</given-names></name> (<year>1997</year>) <article-title>Brunswikian and Thurstonian origins of bias in probability assessment: On the interpretation of stochastic components of judgment</article-title>. <source>J Behav Decis Making</source> <volume>10</volume>: <fpage>189</fpage>–<lpage>209</lpage>.</mixed-citation>
</ref>
<ref id="pone.0096511-Lichtenstein1"><label>17</label>
<mixed-citation publication-type="journal" xlink:type="simple"><name name-style="western"><surname>Lichtenstein</surname><given-names>S</given-names></name>, <name name-style="western"><surname>Fischhoff</surname><given-names>B</given-names></name> (<year>1977</year>) <article-title>Do those who know more also know more about how much they know?</article-title> <source>Organizational Behavior and Human Decision Processes</source> <volume>20</volume>: <fpage>159</fpage>–<lpage>183</lpage>.</mixed-citation>
</ref>
<ref id="pone.0096511-Moore1"><label>18</label>
<mixed-citation publication-type="journal" xlink:type="simple"><name name-style="western"><surname>Moore</surname><given-names>DA</given-names></name>, <name name-style="western"><surname>Healy</surname><given-names>PJ</given-names></name> (<year>2008</year>) <article-title>The trouble with overconfidence</article-title>. <source>Psychol Rev</source> <volume>115</volume>: <fpage>502</fpage>–<lpage>517</lpage>.</mixed-citation>
</ref>
<ref id="pone.0096511-Green1"><label>19</label>
<mixed-citation publication-type="other" xlink:type="simple">Green DM, Swets JA (1966) Signal detection theory and psychophysics. New York,: Wiley. xi, 455 p. p.</mixed-citation>
</ref>
<ref id="pone.0096511-Kepecs2"><label>20</label>
<mixed-citation publication-type="journal" xlink:type="simple"><name name-style="western"><surname>Kepecs</surname><given-names>A</given-names></name>, <name name-style="western"><surname>Mainen</surname><given-names>ZF</given-names></name> (<year>2012</year>) <article-title>A computational framework for the study of confidence in humans and animals</article-title>. <source>Philos Trans R Soc Lond B Biol Sci</source> <volume>367</volume>: <fpage>1322</fpage>–<lpage>1337</lpage>.</mixed-citation>
</ref>
<ref id="pone.0096511-Wichmann1"><label>21</label>
<mixed-citation publication-type="journal" xlink:type="simple"><name name-style="western"><surname>Wichmann</surname><given-names>FA</given-names></name>, <name name-style="western"><surname>Hill</surname><given-names>NJ</given-names></name> (<year>2001</year>) <article-title>The psychometric function: I. Fitting, sampling, and goodness of fit</article-title>. <source>Percept Psychophys</source> <volume>63</volume>: <fpage>1293</fpage>–<lpage>1313</lpage>.</mixed-citation>
</ref>
<ref id="pone.0096511-Britten1"><label>22</label>
<mixed-citation publication-type="journal" xlink:type="simple"><name name-style="western"><surname>Britten</surname><given-names>KH</given-names></name>, <name name-style="western"><surname>Shadlen</surname><given-names>MN</given-names></name>, <name name-style="western"><surname>Newsome</surname><given-names>WT</given-names></name>, <name name-style="western"><surname>Movshon</surname><given-names>JA</given-names></name> (<year>1992</year>) <article-title>The analysis of visual motion: a comparison of neuronal and psychophysical performance</article-title>. <source>J Neurosci</source> <volume>12</volume>: <fpage>4745</fpage>–<lpage>4765</lpage>.</mixed-citation>
</ref>
<ref id="pone.0096511-Newsome1"><label>23</label>
<mixed-citation publication-type="journal" xlink:type="simple"><name name-style="western"><surname>Newsome</surname><given-names>WT</given-names></name>, <name name-style="western"><surname>Britten</surname><given-names>KH</given-names></name>, <name name-style="western"><surname>Movshon</surname><given-names>JA</given-names></name> (<year>1989</year>) <article-title>Neuronal correlates of a perceptual decision</article-title>. <source>Nature</source> <volume>341</volume>: <fpage>52</fpage>–<lpage>54</lpage>.</mixed-citation>
</ref>
<ref id="pone.0096511-Ratcliff2"><label>24</label>
<mixed-citation publication-type="journal" xlink:type="simple"><name name-style="western"><surname>Ratcliff</surname><given-names>R</given-names></name> (<year>1978</year>) <article-title>Theory of Memory Retrieval</article-title>. <source>Psychol Rev</source> <volume>85</volume>: <fpage>59</fpage>–<lpage>108</lpage>.</mixed-citation>
</ref>
<ref id="pone.0096511-Risken1"><label>25</label>
<mixed-citation publication-type="book" xlink:type="simple">Risken H (1989) The Fokker-Planck equation: methods of solution and applications. Berlin; New York: Springer-Verlag. <volume>xiv</volume> , <volume>472</volume>  p. p.</mixed-citation>
</ref>
<ref id="pone.0096511-Drugowitsch1"><label>26</label>
<mixed-citation publication-type="journal" xlink:type="simple"><name name-style="western"><surname>Drugowitsch</surname><given-names>J</given-names></name>, <name name-style="western"><surname>Moreno-Bote</surname><given-names>R</given-names></name>, <name name-style="western"><surname>Churchland</surname><given-names>AK</given-names></name>, <name name-style="western"><surname>Shadlen</surname><given-names>MN</given-names></name>, <name name-style="western"><surname>Pouget</surname><given-names>A</given-names></name> (<year>2012</year>) <article-title>The Cost of Accumulating Evidence in Perceptual Decision Making</article-title>. <source>J Neurosci</source> <volume>32</volume>: <fpage>3612</fpage>–<lpage>3628</lpage>.</mixed-citation>
</ref>
<ref id="pone.0096511-MorenoBote1"><label>27</label>
<mixed-citation publication-type="journal" xlink:type="simple"><name name-style="western"><surname>Moreno-Bote</surname><given-names>R</given-names></name> (<year>2010</year>) <article-title>Decision confidence and uncertainty in diffusion models with partially correlated neuronal integrators</article-title>. <source>Neural Comput</source> <volume>22</volume>: <fpage>1786</fpage>–<lpage>1811</lpage>.</mixed-citation>
</ref>
<ref id="pone.0096511-Erev1"><label>28</label>
<mixed-citation publication-type="journal" xlink:type="simple"><name name-style="western"><surname>Erev</surname><given-names>I</given-names></name>, <name name-style="western"><surname>Wallsten</surname><given-names>TS</given-names></name>, <name name-style="western"><surname>Budescu</surname><given-names>DV</given-names></name> (<year>1994</year>) <article-title>Simultaneous over- and underconfidence: The role of error in judgment processes</article-title>. <source>Psychol Rev</source> <volume>101</volume>: <fpage>519</fpage>–<lpage>527</lpage>.</mixed-citation>
</ref>
<ref id="pone.0096511-Pfeifer1"><label>29</label>
<mixed-citation publication-type="journal" xlink:type="simple"><name name-style="western"><surname>Pfeifer</surname><given-names>PE</given-names></name> (<year>1994</year>) <article-title>Are we overconfident in the belief that probability forecasters are overconfident?</article-title> <source>Organizational Behavior and Human Decision Processes</source> <volume>58</volume>: <fpage>203</fpage>–<lpage>213</lpage>.</mixed-citation>
</ref>
<ref id="pone.0096511-Merkle1"><label>30</label>
<mixed-citation publication-type="journal" xlink:type="simple"><name name-style="western"><surname>Merkle</surname><given-names>EC</given-names></name> (<year>2009</year>) <article-title>The disutility of the hard-easy effect in choice confidence</article-title>. <source>Psychon Bull Rev</source> <volume>16</volume>: <fpage>204</fpage>–<lpage>213</lpage>.</mixed-citation>
</ref>
<ref id="pone.0096511-Baranski1"><label>31</label>
<mixed-citation publication-type="journal" xlink:type="simple"><name name-style="western"><surname>Baranski</surname><given-names>JV</given-names></name>, <name name-style="western"><surname>Petrusic</surname><given-names>WM</given-names></name> (<year>1999</year>) <article-title>Realism of confidence in sensory discrimination</article-title>. <source>Percept Psychophys</source> <volume>61</volume>: <fpage>1369</fpage>–<lpage>1383</lpage>.</mixed-citation>
</ref>
<ref id="pone.0096511-Ferrell1"><label>32</label>
<mixed-citation publication-type="journal" xlink:type="simple"><name name-style="western"><surname>Ferrell</surname><given-names>WR</given-names></name> (<year>1995</year>) <article-title>A model for realism of confidence judgments: implications for underconfidence in sensory discrimination</article-title>. <source>Percept Psychophys</source> <volume>57</volume>: <fpage>246</fpage>–<lpage>254; discussion 255–249</lpage>.</mixed-citation>
</ref>
<ref id="pone.0096511-Ferrell2"><label>33</label>
<mixed-citation publication-type="journal" xlink:type="simple"><name name-style="western"><surname>Ferrell</surname><given-names>WR</given-names></name>, <name name-style="western"><surname>McGoey</surname><given-names>PJ</given-names></name> (<year>1989</year>) <article-title>A model of calibration for subjective probabilities</article-title>. <source>Organizational Behavior and Human Decision Processes</source> <volume>26</volume>: <fpage>32</fpage>–<lpage>53</lpage>.</mixed-citation>
</ref>
<ref id="pone.0096511-Merkle2"><label>34</label>
<mixed-citation publication-type="journal" xlink:type="simple"><name name-style="western"><surname>Merkle</surname><given-names>EC</given-names></name>, <name name-style="western"><surname>Van Zandt</surname><given-names>T</given-names></name> (<year>2006</year>) <article-title>An application of the poisson race model to confidence calibration</article-title>. <source>J Exp Psychol Gen</source> <volume>135</volume>: <fpage>391</fpage>–<lpage>408</lpage>.</mixed-citation>
</ref>
<ref id="pone.0096511-Wald1"><label>35</label>
<mixed-citation publication-type="book" xlink:type="simple">Wald A (1947) Sequential analysis. New York, London: J. Wiley &amp; sons, Chapman &amp; Hall. <volume>xii</volume>: , 212 p. p.</mixed-citation>
</ref>
<ref id="pone.0096511-Wald2"><label>36</label>
<mixed-citation publication-type="journal" xlink:type="simple"><name name-style="western"><surname>Wald</surname><given-names>A</given-names></name>, <name name-style="western"><surname>Wolfowitz</surname><given-names>J</given-names></name> (<year>1948</year>) <article-title>Optimum Character of the Sequential Probability Ratio Test</article-title>. <source>The Annals of Mathematical Statistics</source> <volume>19</volume>: <fpage>326</fpage>–<lpage>339</lpage>.</mixed-citation>
</ref>
<ref id="pone.0096511-Fishman1"><label>37</label>
<mixed-citation publication-type="journal" xlink:type="simple"><name name-style="western"><surname>Fishman</surname><given-names>MM</given-names></name> (<year>1987</year>) <article-title>Average duration of asymptotically optimal multialternative sequential proceure for recognition of processes</article-title>. <source>Soviet Journal of Communication and Technology of Electronics</source> <volume>30</volume>: <fpage>2541</fpage>–<lpage>2548</lpage>.</mixed-citation>
</ref>
<ref id="pone.0096511-Baum1"><label>38</label>
<mixed-citation publication-type="journal" xlink:type="simple"><name name-style="western"><surname>Baum</surname><given-names>CW</given-names></name>, <name name-style="western"><surname>Veeravalli</surname><given-names>VV</given-names></name> (<year>1994</year>) <article-title>A Sequential Procedure for Multihypothesis Testing</article-title>. <source>Ieee T Inform Theory</source> <volume>40</volume>: <fpage>1994</fpage>–<lpage>2007</lpage>.</mixed-citation>
</ref>
<ref id="pone.0096511-Lorden1"><label>39</label>
<mixed-citation publication-type="journal" xlink:type="simple"><name name-style="western"><surname>Lorden</surname><given-names>G</given-names></name> (<year>1977</year>) <article-title>Nearly-Optimal Sequential Tests for Finitely Many Parameter Values</article-title>. <source>Annals of Statistics</source> <volume>5</volume>: <fpage>1</fpage>–<lpage>21</lpage>.</mixed-citation>
</ref>
<ref id="pone.0096511-Dragalin1"><label>40</label>
<mixed-citation publication-type="journal" xlink:type="simple"><name name-style="western"><surname>Dragalin</surname><given-names>VP</given-names></name>, <name name-style="western"><surname>Tartakovsky</surname><given-names>AG</given-names></name>, <name name-style="western"><surname>Veeravalli</surname><given-names>VV</given-names></name> (<year>1999</year>) <article-title>Multihypothesis sequential probability ratio tests - Part I: Asymptotic optimality</article-title>. <source>Ieee T Inform Theory</source> <volume>45</volume>: <fpage>2448</fpage>–<lpage>2461</lpage>.</mixed-citation>
</ref>
<ref id="pone.0096511-Beck2"><label>41</label>
<mixed-citation publication-type="journal" xlink:type="simple"><name name-style="western"><surname>Beck</surname><given-names>JM</given-names></name>, <name name-style="western"><surname>Ma</surname><given-names>WJ</given-names></name>, <name name-style="western"><surname>Pitkow</surname><given-names>X</given-names></name>, <name name-style="western"><surname>Latham</surname><given-names>PE</given-names></name>, <name name-style="western"><surname>Pouget</surname><given-names>A</given-names></name> (<year>2012</year>) <article-title>Not noisy, just wrong: the role of suboptimal inference in behavioral variability</article-title>. <source>Neuron</source> <volume>74</volume>: <fpage>30</fpage>–<lpage>39</lpage>.</mixed-citation>
</ref>
<ref id="pone.0096511-Kahneman1"><label>42</label>
<mixed-citation publication-type="book" xlink:type="simple">Kahneman D, Slovic P, Tversky A (1982) Judgment under uncertainty: heuristics and biases. Cambridge: Cambridge University Press. <volume>xiii</volume>: , 555 p. p.</mixed-citation>
</ref>
<ref id="pone.0096511-Zylberberg1"><label>43</label>
<mixed-citation publication-type="journal" xlink:type="simple"><name name-style="western"><surname>Zylberberg</surname><given-names>A</given-names></name>, <name name-style="western"><surname>Barttfeld</surname><given-names>P</given-names></name>, <name name-style="western"><surname>Sigman</surname><given-names>M</given-names></name> (<year>2012</year>) <article-title>The construction of confidence in a perceptual decision</article-title>. <source>Front Integr Neurosci</source> <volume>6</volume>: <fpage>79</fpage>.</mixed-citation>
</ref>
<ref id="pone.0096511-Battaglia1"><label>44</label>
<mixed-citation publication-type="journal" xlink:type="simple"><name name-style="western"><surname>Battaglia</surname><given-names>PW</given-names></name>, <name name-style="western"><surname>Jacobs</surname><given-names>RA</given-names></name>, <name name-style="western"><surname>Aslin</surname><given-names>RN</given-names></name> (<year>2003</year>) <article-title>Bayesian integration of visual and auditory signals for spatial localization</article-title>. <source>J Opt Soc Am A Opt Image Sci Vis</source> <volume>20</volume>: <fpage>1391</fpage>–<lpage>1397</lpage>.</mixed-citation>
</ref>
<ref id="pone.0096511-Galvin1"><label>45</label>
<mixed-citation publication-type="journal" xlink:type="simple"><name name-style="western"><surname>Galvin</surname><given-names>SJ</given-names></name>, <name name-style="western"><surname>Podd</surname><given-names>JV</given-names></name>, <name name-style="western"><surname>Drga</surname><given-names>V</given-names></name>, <name name-style="western"><surname>Whitmore</surname><given-names>J</given-names></name> (<year>2003</year>) <article-title>Type 2 tasks in the theory of signal detectability: discrimination between correct and incorrect decisions</article-title>. <source>Psychon Bull Rev</source> <volume>10</volume>: <fpage>843</fpage>–<lpage>876</lpage>.</mixed-citation>
</ref>
<ref id="pone.0096511-Maniscalco1"><label>46</label>
<mixed-citation publication-type="journal" xlink:type="simple"><name name-style="western"><surname>Maniscalco</surname><given-names>B</given-names></name>, <name name-style="western"><surname>Lau</surname><given-names>H</given-names></name> (<year>2012</year>) <article-title>A signal detection theoretic approach for estimating metacognitive sensitivity from confidence ratings</article-title>. <source>Conscious Cogn</source> <volume>21</volume>: <fpage>422</fpage>–<lpage>430</lpage>.</mixed-citation>
</ref>
<ref id="pone.0096511-Smith2"><label>47</label>
<mixed-citation publication-type="journal" xlink:type="simple"><name name-style="western"><surname>Smith</surname><given-names>PL</given-names></name> (<year>2000</year>) <article-title>Stochastic Dynamic Models of Response Time and Accuracy: A Foundational Primer</article-title>. <source>J Math Psychol</source> <volume>44</volume>: <fpage>408</fpage>–<lpage>463</lpage>.</mixed-citation>
</ref>
<ref id="pone.0096511-Cox1"><label>48</label>
<mixed-citation publication-type="book" xlink:type="simple">Cox DR, Miller HD (1965) The theory of stochastic processes. New York: Wiley. <volume>x</volume>: , 398 p. p.</mixed-citation>
</ref>
<ref id="pone.0096511-Palmer1"><label>49</label>
<mixed-citation publication-type="journal" xlink:type="simple"><name name-style="western"><surname>Palmer</surname><given-names>J</given-names></name>, <name name-style="western"><surname>Huk</surname><given-names>AC</given-names></name>, <name name-style="western"><surname>Shadlen</surname><given-names>MN</given-names></name> (<year>2005</year>) <article-title>The effect of stimulus strength on the speed and accuracy of a perceptual decision</article-title>. <source>J Vis</source> <volume>5</volume>: <fpage>376</fpage>–<lpage>404</lpage>.</mixed-citation>
</ref>
</ref-list></back>
</article>