Secondary motor integration as a final arbiter in sensorimotor decision-making.

Sensorimotor decision-making is believed to involve a process of accumulating sensory evidence over time. While current theories posit a single accumulation process prior to planning an overt motor response, here, we propose an active role of motor processes in decision formation via a secondary leaky motor accumulation stage. The motor leak adapts the "memory" with which this secondary accumulator reintegrates the primary accumulated sensory evidence, thus adjusting the temporal smoothing in the motor evidence and, correspondingly, the lag between the primary and motor accumulators. We compare this framework against different single accumulator variants using formal model comparison, fitting choice, and response times in a task where human observers made categorical decisions about a noisy sequence of images, under different speed-accuracy trade-off instructions. We show that, rather than boundary adjustments (controlling the amount of evidence accumulated for decision commitment), adjustment of the leak in the secondary motor accumulator provides the better description of behavior across conditions. Importantly, we derive neural correlates of these 2 integration processes from electroencephalography data recorded during the same task and show that these neural correlates adhere to the neural response profiles predicted by the model. This framework thus provides a neurobiologically plausible description of sensorimotor decision-making that captures emerging evidence of the active role of motor processes in choice behavior.

Metacognitive evaluation of postdecisional perceptual representations.

Perceptual confidence is thought to arise from metacognitive processes that evaluate the underlying perceptual decision evidence. We investigated whether metacognitive access to perceptual evidence is constrained by the hierarchical organization of visual cortex, where high-level representations tend to be more readily available for explicit scrutiny. We found that the ability of human observers to evaluate their confidence did depend on whether they performed a high-level or low-level task on the same stimuli, but was also affected by manipulations that occurred long after the perceptual decision. Confidence in low-level perceptual decisions degraded with more time between the decision and the response cue, especially when backward masking was present. Confidence in high-level tasks was immune to backward masking and benefitted from additional time. These results can be explained by a model assuming confidence heavily relies on postdecisional internal representations of visual stimuli that degrade over time, where high-level representations are more persistent.

Absolute and relative blindsight.

The concept of relative blindsight, referring to a difference in conscious awareness between conditions otherwise matched for performance, was introduced by Lau and Passingham (2006) as a way of identifying the neural correlates of consciousness (NCC) in fMRI experiments. By analogy, absolute blindsight refers to a difference between performance and awareness regardless of whether it is possible to match performance across conditions. Here, we address the question of whether relative and absolute blindsight in normal observers can be accounted for by response bias. In our replication of Lau and Passingham's experiment, the relative blindsight effect was abolished when performance was assessed by means of a bias-free 2AFC task or when the criterion for awareness was varied. Furthermore, there was no evidence of either relative or absolute blindsight when both performance and awareness were assessed with bias-free measures derived from confidence ratings using signal detection theory. This suggests that both relative and absolute blindsight in normal observers amount to no more than variations in response bias in the assessment of performance and awareness. Consideration of the properties of psychometric functions reveals a number of ways in which relative and absolute blindsight could arise trivially and elucidates a basis for the distinction between Type 1 and Type 2 blindsight.

Distinct basal ganglia contributions to learning from implicit and explicit value signals in perceptual decision-making.

Metacognitive evaluations of confidence provide an estimate of decision accuracy that could guide learning in the absence of explicit feedback. We examine how humans might learn from this implicit feedback in direct comparison with that of explicit feedback, using simultaneous EEG-fMRI. Participants performed a motion direction discrimination task where stimulus difficulty was increased to maintain performance, with intermixed explicit- and no-feedback trials. We isolate single-trial estimates of post-decision confidence using EEG decoding, and find these neural signatures re-emerge at the time of feedback together with separable signatures of explicit feedback. We identified these signatures of implicit versus explicit feedback along a dorsal-ventral gradient in the striatum, a finding uniquely enabled by an EEG-fMRI fusion. These two signals appear to integrate into an aggregate representation in the external globus pallidus, which could broadcast updates to improve cortical decision processing via the thalamus and insular cortex, irrespective of the source of feedback.

Confidence at the limits of human nested cognition.

Metacognition is the ability to weigh the quality of our own cognition, such as the confidence that our perceptual decisions are correct. Here we ask whether metacognitive performance can itself be evaluated or else metacognition is the ultimate reflective human faculty. Building upon a classic visual perception task, we show that human observers are able to produce nested, above-chance judgements on the quality of their decisions at least up to the fourth order (i.e. meta-meta-meta-cognition). A computational model can account for this nested cognitive ability if evidence has a high-resolution representation, and if there are two kinds of noise, including recursive evidence degradation. The existence of fourth-order sensitivity suggests that the neural mechanisms responsible for second-order metacognition can be flexibly generalized to evaluate any cognitive process, including metacognitive evaluations themselves. We define the theoretical and practical limits of nested cognition and discuss how this approach paves the way for a better understanding of human self-regulation.

Consensus Goals in the Field of Visual Metacognition.

Despite the tangible progress in psychological and cognitive sciences over the last several years, these disciplines still trail other more mature sciences in identifying the most important questions that need to be solved. Reaching such consensus could lead to greater synergy across different laboratories, faster progress, and increased focus on solving important problems rather than pursuing isolated, niche efforts. Here, 26 researchers from the field of visual metacognition reached consensus on four long-term and two medium-term common goals. We describe the process that we followed, the goals themselves, and our plans for accomplishing these goals. If this effort proves successful within the next few years, such consensus building around common goals could be adopted more widely in psychological science.

Separable neural signatures of confidence during perceptual decisions.

Perceptual confidence is an evaluation of the validity of perceptual decisions. While there is behavioural evidence that confidence evaluation differs from perceptual decision-making, disentangling these two processes remains a challenge at the neural level. Here, we examined the electrical brain activity of human participants in a protracted perceptual decision-making task where observers tend to commit to perceptual decisions early whilst continuing to monitor sensory evidence for evaluating confidence. Premature decision commitments were revealed by patterns of spectral power overlying motor cortex, followed by an attenuation of the neural representation of perceptual decision evidence. A distinct neural representation was associated with the computation of confidence, with sources localised in the superior parietal and orbitofrontal cortices. In agreement with a dissociation between perception and confidence, these neural resources were recruited even after observers committed to their perceptual decisions, and thus delineate an integral neural circuit for evaluating perceptual decision confidence.

Confidence controls perceptual evidence accumulation.

Perceptual decisions are accompanied by feelings of confidence that reflect the likelihood that the decision was correct. Here we aim to clarify the relationship between perception and confidence by studying the same perceptual task across three different confidence contexts. Human observers were asked to categorize the source of sequentially presented visual stimuli. Each additional stimulus provided evidence for making more accurate perceptual decisions, and better confidence judgements. We show that observers' ability to set appropriate evidence accumulation bounds for perceptual decisions is strongly predictive of their ability to make accurate confidence judgements. When observers were not permitted to control their exposure to evidence, they imposed covert bounds on their perceptual decisions but not on their confidence decisions. This partial dissociation between decision processes is reflected in behaviour and pupil dilation. Together, these findings suggest a confidence-regulated accumulation-to-bound process that controls perceptual decision-making even in the absence of explicit speed-accuracy trade-offs.

How wide is the cone of direct gaze?

The cone of direct gaze refers to the range of gaze deviations an observer accepts as looking directly at them. Previous experiments have calculated the width of the cone of direct gaze using the gaze deviations actually presented to the observer, however, there is considerable evidence that observers actually perceive gaze to be systematically more deviated than actually presented. Here, we examine the width of the cone of direct gaze in units of perceived gaze deviation. In doing so, we are able to disambiguate differences in width both within and between observers that are due to differences in their perception of gaze and due to differences in what observers consider to be looking at them. We suggest that this line of inquiry can offer further insight into the perception of gaze direction, and how this perception may differ in clinical populations.

Task Dependent Effects of Head Orientation on Perceived Gaze Direction.

The perception of gaze direction involves the integration of a number of sensory cues exterior to the eye-region. The orientation of the head is one such cue, which has an overall repulsive effect on the perceived direction of gaze. However, in a recent experiment, we found the measured effect of head orientation on perceived gaze direction differed within subjects, depending on whether a single- or two-interval task design was employed. This suggests a potential difference in the way the orientation of the head is integrated into the perception of gaze direction across tasks. Four experiments were conducted to investigate this difference. The first two experiments showed that the difference was not the result of some interaction between stimuli in the two-interval task, but rather, a difference between the types of judgment being made across tasks, where observers were making a directional (left/right) judgment in the single-interval task, and a non-directional (direct/indirect gaze) judgment in the two-interval task. A third experiment showed that this difference does not arise from observers utilizing a non-directional cue to direct gaze (the circularity of the pupil/iris) in making their non-directional judgments. The fourth experiment showed no substantial differences in the duration of evidence accumulation and processing between judgments, suggesting that observers are not integrating different sensory information across tasks. Together these experiments show that the sensory information from head orientation is flexibly weighted in the perception of gaze direction, and that the purpose of the observer, in sampling gaze information, can influence the consequent perception of gaze direction.

Visual processing: conscious until proven otherwise.

Unconscious perception, or perception without awareness, describes a situation where an observer's behaviour is influenced by a stimulus of which they have no phenomenal awareness. Perception without awareness is often claimed on the basis of a difference in thresholds for tasks that do and do not require awareness, for example, detecting the stimulus (requiring awareness) and making accurate judgements about the stimulus (based on unconscious processing). Although a difference in thresholds would be expected if perceptual evidence were processed without awareness, such a difference does not necessitate that this is actually occurring: a difference in thresholds can also arise from response bias, or through task differences. Here we ask instead whether the pattern of performance could be obtained if the observer were aware of the evidence used in making their decisions. A backwards masking paradigm was designed using digits as target stimuli, with difficulty controlled by the time between target and mask. Performance was measured over three tasks: detection, graphic discrimination and semantic discrimination. Despite finding significant differences in thresholds measured using proportion correct, and in observer sensitivity, modelling suggests that these differences were not the result of perception without awareness. That is, the observer was not relying solely on unconscious information to make decisions.

Improving face identification with specialist teams.

People vary in their ability to identify faces, and this variability is relatively stable across repeated testing. This suggests that recruiting high performers can improve identity verification accuracy in applied settings. Here, we report the first systematic study to evaluate real-world benefits of selecting high performers based on performance in standardized face identification tests. We simulated a recruitment process for a specialist team tasked with detecting fraudulent passport applications. University students (n = 114) completed a battery of screening tests followed by a real-world face identification task that is performed routinely when issuing identity documents. Consistent with previous work, individual differences in the real-world task were relatively stable across repeated tests taken 1 week apart (r = 0.6), and accuracy scores on screening tests and the real-world task were moderately correlated. Nevertheless, performance gains achieved by selecting groups based on screening tests were surprisingly small, leading to a 7% improvement in accuracy. Statistically aggregating decisions across individuals-using a 'wisdom of crowds' approach-led to more substantial gains than selection alone. Finally, controlling for individual accuracy of team members, the performance of a team in one test predicted their performance in a subsequent test, suggesting that a 'good team' is not only defined by the individual accuracy of team members. Overall, these results underline the need to use a combination of approaches to improve face identification performance in professional settings.

A bias-minimising measure of the influence of head orientation on perceived gaze direction.

The orientation of the head is an important cue for gaze direction, and its role has been explained in a dual route model. The model incorporates both an attractive and a repulsive effect of head orientation, which act to support accurate gaze perception across large changes in natural stimuli. However, in all previous studies of which we are aware, measurements of the influence of head orientation on perceived gaze direction were obtained using a single-interval methodology, which may have been affected by response bias. Here we compare the single-interval methodology with a two-interval (bias-minimising) design. We find that although measures obtained using the two-interval design showed a stronger attractive effect of head orientation than previous studies, the influence of head orientation on perceived gaze direction still represents a genuine perceptual effect. Measurements obtained using the two-interval design were also shown to be more stable across sessions one week apart. These findings suggest the two-interval design should be used in future experiments, especially if comparing groups who may systematically differ in their biases, such as patients with schizophrenia or autism.

Detecting and identifying offset gaze.

A number of experiments have demonstrated that observers can accurately identify stimuli that they fail to detect (Rollman and Nachmias, 1972; Harris and Fahle, 1995; Allik et al. 1982, 2014). Using a 2x2AFC double judgements procedure, we demonstrated an analogous pattern of performance in making judgements about the direction of eye gaze. Participants were shown two faces in succession: one with direct gaze and one with gaze offset to the left or right. We found that they could identify the direction of gaze offset (left/right) better than they could detect which face contained the offset gaze. A simple Thurstonian model, under which the detection judgement is shown to be more computationally complex, was found to explain the empirical data. A further experiment incorporated metacognitive ratings into the double judgements procedure to measure observers' metacognitive awareness (Meta-d') across the two judgements and to assess whether observers were aware of the evidence for offset gaze when detection performance was at and below threshold. Results suggest that metacognitive awareness is tied to performance, with approximately equal Meta-d' across the two judgements, when sensitivity is taken into account. These results show that both performance and metacognitive awareness rely not only on the strength of sensory evidence but also on the computational complexity of the decision, which determines the relative distance of that evidence from the decision axes.

Depression Reduces Accuracy While Parkinsonism Slows Response Time for Processing Positive Feedback in Patients with Parkinson's Disease with Comorbid Major Depressive Disorder Tested on a Probabilistic Category-Learning Task.

Major depressive disorder (MDD) is the most common non-motor manifestation of Parkinson's disease (PD) affecting 50% of patients. However, little is known about the cognitive correlates of MDD in PD. Using a computer-based cognitive task that dissociates learning from positive and negative feedback, we tested four groups of subjects: (1) patients with PD with comorbid MDD, (2) patients with PD without comorbid MDD, (3) matched patients with MDD alone (without PD), and (4) matched healthy control subjects. Furthermore, we used a mathematical model of decision-making to fit both choice and response time data, allowing us to detect and characterize differences between the groups that are not revealed by cognitive results. The groups did not differ in learning accuracy from negative feedback, but the MDD groups (PD patients with MDD and patients with MDD alone) exhibited a selective impairment in learning accuracy from positive feedback when compared to the non-MDD groups (PD patients without MDD and healthy subjects). However, response time in positive feedback trials in the PD groups (both with and without MDD) was significantly slower than the non-PD groups (MDD and healthy groups). While faster response time usually correlates with poor learning accuracy, it was paradoxical in PD groups, with PD patients with MDD having impaired learning accuracy and PD patients without MDD having intact learning accuracy. Mathematical modeling showed that both MDD groups (PD with MDD and MDD alone) were significantly slower than non-MDD groups in the rate of accumulation of information for stimuli trained by positive feedback, which can lead to lower response accuracy. Conversely, modeling revealed that both PD groups (PD with MDD and PD alone) required more evidence than other groups to make responses, thus leading to slower response times. These results suggest that PD patients with MDD exhibit cognitive profiles with mixed traits characteristic of both MDD and PD, furthering our understanding of both PD and MDD and their often-complex comorbidity. To the best of our knowledge, this is the first study to examine feedback-based learning in PD with MDD while controlling for the effects of PD and MDD.

Probabilistic reward- and punishment-based learning in opioid addiction: Experimental and computational data.

Addiction is the continuation of a habit in spite of negative consequences. A vast literature gives evidence that this poor decision-making behavior in individuals addicted to drugs also generalizes to laboratory decision making tasks, suggesting that the impairment in decision-making is not limited to decisions about taking drugs. In the current experiment, opioid-addicted individuals and matched controls with no history of illicit drug use were administered a probabilistic classification task that embeds both reward-based and punishment-based learning trials, and a computational model of decision making was applied to understand the mechanisms describing individuals' performance on the task. Although behavioral results showed that opioid-addicted individuals performed as well as controls on both reward- and punishment-based learning, the modeling results suggested subtle differences in how decisions were made between the two groups. Specifically, the opioid-addicted group showed decreased tendency to repeat prior responses, meaning that they were more likely to "chase reward" when expectancies were violated, whereas controls were more likely to stick with a previously-successful response rule, despite occasional expectancy violations. This tendency to chase short-term reward, potentially at the expense of developing rules that maximize reward over the long term, may be a contributing factor to opioid addiction. Further work is indicated to better understand whether this tendency arises as a result of brain changes in the wake of continued opioid use/abuse, or might be a pre-existing factor that may contribute to risk for addiction.

Drift diffusion model of reward and punishment learning in schizophrenia: Modeling and experimental data.

In this study, we tested reward- and punishment learning performance using a probabilistic classification learning task in patients with schizophrenia (n=37) and healthy controls (n=48). We also fit subjects' data using a Drift Diffusion Model (DDM) of simple decisions to investigate which components of the decision process differ between patients and controls. Modeling results show between-group differences in multiple components of the decision process. Specifically, patients had slower motor/encoding time, higher response caution (favoring accuracy over speed), and a deficit in classification learning for punishment, but not reward, trials. The results suggest that patients with schizophrenia adopt a compensatory strategy of favoring accuracy over speed to improve performance, yet still show signs of a deficit in learning based on negative feedback. Our data highlights the importance of applying fitting models (particularly drift diffusion models) to behavioral data. The implications of these findings are discussed relative to theories of schizophrenia and cognitive processing.